Current Problems of Agrarian Industry in Ukraine

CURRENT PROBLEMS OF AGRARIAN INDUSTRY IN UKRAINE

Accent Graphics Communications & Publishing

Vancouver 2019 Reviewers: Gritsan Y. I. – Doctor of Science (Biology), Professor of the Department of Ecology and Environmental Protection, Dnipro State Agrarian and Economic University; Lykholat Y. V. – Doctor of Science (Biology), Professor, Head of the Department of Physiology and Introduction of Plants, Oles Honchar Dnipro National University. Skliarov P. M. – Doctor of Science (Veterinary medicine), Professor of the Department of Surgery and Obstetrics of Farm Animals, Dnipro State Agrarian and Economic University

Approved by the Academic Council of Dnipro State Agrarian and Economic University (protocol № 9 from 27.06.2019) Current problems of agrarian industry in Ukraine. Accent Graphics Communications & Publishing, Vancouver, Canada, 2019. – 228 p. ISBN 978-1-77192-487-0 DOI: http://doi.org/10.29013/NMZazharska.CPAIU.228.2019

The monograph is presented in four parts. The first part is devoted to the experimental and theoretical substantiation of the criteria for safety and quality assessment of goat's milk. Parameters of subclinical mastitis in goats, comparison of methods efficiency for determination of somatic cell count in goat milk, monitoring studies of goat’s and cow’s milk in France and Ukraine, effect of exogenous and endogenous factors on the quality and safety of goat milk are described. The second and third parts are devoted to Coleoptera pests of stored food supplies and field crops. The forth part includes characteristics of Poaceae family members in the steppe zone of Ukraine as the main objects of farm animals feeding. Ecological characteristics of the species according to the Belgard Ekomorph System and their geographical analysis were presented. Sozological and feeding value of the species were described.

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Chapter 1. Assessment of safety and quality of goat’s milk

N. M. Zazharska Dnipro State Agrarian and Economic University

The criteria for safety and quality assessment of goat’s milk 1

According to the European regulation 853/2004 only microbiological criteria have been put forward for goat milk: bacterial contamination at a temperature of 30 °C is ≤ 1 500 × 103 CFU/cm3; in milk intended for the production of products without heat treatment the content of microorganisms – ≤ 500×103 CFU/cm3. It is substantiated that the Na- tional Standards of Ukraine (DSTU) “Goat’s milk. Raw. Specifications: DSTU7006: 2009” does not meet the international requirements for goat milk (Zazharska, 2018). According to the requirements of existing normative document DSTU7006:2009 the number of mesophilic aerobic and extra-anaerobic microorganisms of milk of the second grade (the extreme limit) corresponds to the best milk in accordance with the European regulations. According to Ukraine requirements it is permitted somatic cell count to 500 × 103 cells/cm3, bacterial contamination ≤ 100 × 103 colony forming units (CFU)/cm3 for the highest quality goat milk. In most European countries and the USA, the best goat milk is considered with somatic cell count ≤ 1 million/cm3, bacterial contamination ≤ 500 ×103 CFU/cm3. Consequently, the requirements for the indexes of bacterial contamination and somatic cell count in the Ukrainian normative document are very strict. Indicators of density and acidity of goat milk are not regulated in EU countries, but regulated in Ukrainian standard.

1 Zazharska N. M. (2019). Assessment of safety and quality of goat’s milk. In: Current problems of agrarian industry in Ukraine. Accent Graphics Communications & Publishing, Vancouver, Canada. – P. 3–33. Doi:10.15421/511902 4 Chapter 1. Assessment of safety and quality of goat’s milk

The aim was to scientifically substantiate the requirements of the Ukrainian standard for goat milk, especially for indicators of acidity, density, somatic cell count and bacterial contamination. 836 samples of goat milk were taken from farms of Dnipropetrovs’k and Transcarpathian oblasts during 2013–2017 years. Physico-chemical parameters (acidity, density, indicators of fat and protein) of goat’s milk were determined using the ultrasound analyzer “Ekomilk”. The somatic cell count was determined by viscosimetric method and flow cytometry. According to our research the indicators of the acidity in goat milk vary from 14 °T to 27 °T, of the density – from 25.6 °A to 35.4 °A. The smallest somatic cell count in the goat milk was observed in the autumn 265 ± 41 × 103 cells/cm3, the highest – in the winter – 451 ± ± 46 × 103 cells/cm3 (p < 0.05) (viscosimetric method). The smallest somatic cell count throughout the life of the animal was observed in the milk of goats of the first lactation – 712 ± 174 × 103 cells/cm3. In goats from the second to the fourth lactation, somatic cell count was recorded at the level of 880–1092 × 103 cells/cm3 (flow cytometry method). Indicators of fat and protein in goat milk are drastically reduced in the summer, therefore the requirements of the Ukrainian standard of 3.3 and 3.0% respectively are not substantiated. According to the obtained data it was determined that the somatic cell count in goat milk exceeds the index from DSTU7006: 2009 “Goat’s milk. Raw. Specifications: DSTU7006: 2009”, twice or more. It was found that the existing standard does not meet the international requirements for goat milk based on the main indicators (bacterial contamination, somatic cells, acidity and density). The ineffectiveness of the criteria of density and acidity for goat milk assessment was substantiated because of their insignificance. It was proposed to exclude the indexes of density and acidity of goat’s milk from DSTU, to decrease the requirements for the fat and protein content, and to align the requirements for the somatic cell count and bacterial contamination in accordance with Euro- pean standards. N. M. Zazharska 5 Parameters of subclinical mastitis in goats

The universal definition of a somatic cell count threshold to dis- tinguish between healthy and sick udder halves in goats does not exist yet. The aim of this work was to establish the possible diagnostic parameters of subclinical mastitis in goats. 27 samples milk of goats were researched. The main parameters of milk were analyzed by means of ultra- sonic analyzer of milk of “Ekomilk type MILKANA KAM 98–2a” (Bulgary), the somatic cell count was determined at viscometric analyzer “Somatos-M” (Russia), milk films were stained with pironin Y and May-Grünwald methods. Chloride ion content in milk was determined by titrimetric method. Chlorine-sugar number was counted, settling test and mastidin test were also conducted. The data were analysed in Statisica 6.0 (StatSoft Inc., USA). The data in the tables are presented as x ± SE (x ± standard error). The differences between the values in groups were determined using Tukey test, where the differences were considered significant at P < 0.05 (with taking into account the Bonferroni correction). The samples were divided into 3 groups after determination of biochemical parameters, according to the chloride content in the milk: group I – < 250 mg%; group II – 250–300 mg%; group III – > >300 mg% (tab. 1). The indexes of chloride content were significantly different between three groups of samples (P < 0.001). The index of fat content increased by 0.4% in goat’s milk with a chloride concentration > 300 mg%, protein – by 0.24%, lactose – by 0.28%, milk solids – by 0.66%, pH – by 2.8%, the freezing point decreased by 7.2% in comparison to I group of milk samples (with a chloride content < 250 mg%), but it was not statistical difference. In the group of samples with a chloride content > 300 mg%, the somatic cell count increased by 3.2–5.7 times compared to the group with chloride concentration < 250 mg%, depending on the method of study (P < 0.05 and P < 0.001 accordingly). The parameters of somatic cell count of the second group (with a chloride content 6 Chapter 1. Assessment of safety and quality of goat’s milk

250–300 mg%), were 2.1–3.8 times higher compared with the first group (P < 0.05 and P < 0.01 respectively). The chlorine-sugar figure in samples of milk of healthy goats (with a chloride content < 250 mg%) is average 5 (from 4.1 to 5.9). The chlorine-sugar figure is 7.2 (from 6.5 to 7.9) in milk samples of III group (with a chloride content > 300 mg%). Table 1. – Parameters of goat milk depending on the content of chlorides (x ± SE) Milk sample groups according Parameter to the content of chlorides, mg% І, n = 8 ІІ, n =13 ІІІ, n = 6 Chloride content, mg % 223.7 ± 4,7a 270.4 ± 3,8b 344.4 ± 14,8c somatic cell count, × 439 ± 159a 1672 ± 292b 2500 ± 316b 103 cells/ml: “Somatos” May-Grünwald 634 ± 169a 1569 ± 323b 2149 ± 560b pironin Y 703 ± 213a 1484 ± 276b 2273 ± 539b Fat,% 4.35 ± 0,74 4.28 ± 0,47 4.75 ± 0,62 Dry non-fat milk sol- 8.20 ± 0.27 8.13 ± 0.10 8.86 ± 0.34 ids,% Density, ºА 27.2 ± 1.4 27.0 ± 0.4 29.6 ± 1.7 Protein,% 3.06 ± 0.09 3.02 ± 0.04 3.30 ± 0.12 –0.540 ± –0.533 ± –0.579 ± Freezing point, ºС ± 0.015 ± 0.006 ± 0.023 Lactose,% 4.52 ± 0.16 4.48 ± 0.05 4.80 ± 0.24 Conductivity, mS/сm 4.73 ± 0.26a 5.35 ± 0.13b 6.21 ± 0.46b pH 6.71 ± 0.04 6.67 ± 0.04 6.90 ± 0.16 Chlorine-sugar figure 5.00 ± 0.23a 6.05 ± 0.14b 7.21 ± 0.27c Bacterial contamina- 1.1 ± 0.24 × 9.0 ± 4.9 × 1.6 ± 0.5 × tion, CFU/cm3 ×105 ×105 ×106 Mastitis pathogens – – + Mastidin test – + + Settling test – – + Note: different letters within the line correspond to the selections which had significant differences between one another according to the results of Tukey’s test (P < 0.05) with Bonferroni correction N. M. Zazharska 7

The chlorine-sugar figure in the milk samples of the I group was significantly less than in the II and III groups by 20.9% and 44% respectively (P < 0.001). This parameter in the goat milk of the III group is more than in the II group by 19% (P < 0.01). The electrical conductivity in the samples of the I group was less than in the II by 13.1% and than in the III – by 31.3%, (P < 0.05). Н. Schüppel & М. Schwope determined the average electrical conductivity in goat milk 6.6 ± 0.5 mS/сm (Schüppel & Schwope, 1999). Unfortunately, these indicators do not coincide with the results of our research – the electrical conductivity of milk in healthy goats was within the limits 4.73 ± 0.26 mS/сm. However, the absolute threshold for the differentiation of infected and uninfected mammary glands of goats has not yet been found (Barth, 2009). A significant correlation between the electrical conductivity and the somatic cell count that is known in dairy cows does not seem to exist in dairy goats (Park, 1991). Positive settling test was observed in the samples of III group (with a chloride content > 300 mg %). Positive mastidin test with goat’s milk was observed in the samples of II and of III group. Bacterial contamination increased with the growing of the chloride content in goat’s milk, but statistical difference was not been detected. Streptococcus agalactiae was isolated in 2 of 6 milk samples of the III group. It was established that combination of such indexes as the somatic cell count > 2 million/ml, the chloride ion content > 300 mg%, chlorine-sugar figure 7 and above, a positive settling test and mastidin test, can serve as a criterion for detecting subclinical mastitis in goats (Zazharska et al., 2017 b). The study of goat milk on subclinical mastitis and comparison of different methods of determining the somatic cell count was carry out. The herd of goats, German white, Anglo-Nubian and Alpine and local breeds, was studied on the subclinical mastitis twice: in the fall 83 dairy goats were examined, in the spring – 144 animals. The first portions of milk were collected, after which they were examined by mastidin test and Californian mastitis test. Samples of milk were taken to the Laboratory of Food Hygiene at the Department of Parasitology, Veterinary and Sanitary Expertise at the Dnipro State 8 Chapter 1. Assessment of safety and quality of goat’s milk

Agrarian and Economic University. The settling test was conducted there. The determination of somatic cell count in milk was carried out by viscosimetric method. We also made milk films and stained them by the May-Grünwald method (Fig. 1). After that the somatic cell count using microscope was calculat- ed. As a result of bacteriological research of milk on pathogens mastitis Staphylococcus aureus was isolated in autumn. For six months the number of goats with subclinical mastitis decreased from 12 to 8%, by improving the control of the udder health of the animals on the farm. The mastidin test was better than the California mastitis test with goat milk, due to the formation of a tighter clot. From the milk samples which were positive or questionable on mastidin test, 29% samples were found consistent with the requirements of DSTU7006: 2009 “Goat’s milk. Raw. Specifications” for second grade of goat’s milk by viscosimetric method and settling test.

Figure 1. Somatic cells in smears of milk of goat with subclinical mastitis: May-Grünwald stain; bar – 10 μm But by the arbitration method (direct microscopy) the somatic cell count did not meet the requirements of DSTU7006:2009 in all samples. The exact somatic cell count in goat’s milk should be determined only by direct microscopic or fluoroptoelectronic counting. In milk samples, which revealed the largest somatic cell count in milk films (> 20 million/ml), only 2 818 ± 956 × 103 cells/cm3 was determined by viscometric method, which proves the accuracy of the arbitration N. M. Zazharska 9 method. To obtain milk with a low somatic cell count using the viscometric method (< 600 thousand/cm3) for high-quality cheese production a continuous renovation of the herd was recommended because the lowest number of somatic cells in milk observed in goats of the first lactation (primiparous goats) (Zazharska & Rosenko, 2018).

Comparison of methods efficiency for determination of somatic cell count in goat milk

It was compared different methods of determination of somatic cell count in goat milk (Zazharska & Zharko, 2016). Somatic cell count of individual milk samples from 28 goats were analyzed by means of analyzer “Somatos” (viscosimetric method), “SomaCount- FlowCytometer” (flow cytometry) and the by counting of cells in milk films stained with pironin Y, at May-Grünwald and Ro- manovsky – Giemsa methods. There were not found the samples with somatic cells count to 100 × 103 cells/ml while counting cells in milk films stained by any method. According to the research by means of "Soma- CountFlowCytometer" and "Somatos" most of milk smears be-longed to the level of – 101–500 × 103 cells/ml – 35.7 and 50% re-spectively. The biggest part of milk smears – stained by Roma-novsky – Giemsa method – 42.9% and by May-Grünwald method – 39.3% related to the range of 1001–3000 × 103 cells/ml, whereas films, stained with pironin Y and methyl green – 35.7% – to a range of 501–1000 × 103 cells/ml. The greater indexes of somatic cell count in direct counting of cells in goat’s milk smears, stained by any method was determined than using devices. It was proved the accuracy of direct counting method because the distribution ranges of somatic cells was similar between different methods of film staining. The cytoplasm and nuclei of somatic cells are well stained in goat milk films stained by May-Grünwald method (Fig. 2). So, this method is proposed for counting of somatic cells by Prescott and Breed method in goat’s milk. It is proved that the quality of goat's milk films stained by the May-Grünwald method for counting of somatic cells corresponds 10 Chapter 1. Assessment of safety and quality of goat’s milk to the recommended method with pyro-nin Y (Fig. 3), and the cost of dyes is lower by 28.4 times. The Romanovsky – Giemsa method is not quite acceptable for goat's milk films, because we have received higher rates of somatic cell count than other methods of staining because of cytoplasmic particles. Also "shadows" of cells, particles of a paint appear often in milk smears (Fig. 4). So, for the veterinary medicine practice, it is proposed: method of the May-Grünwald for staining of goat's milk films for counting somatic cells by Prescott and Breed method.

Figure 2. Somatic cells in goat's milk smears: May-Grünwald stain; bar – 10 μm

Figure 3. Somatic cells in goat's milk smears (control): pyronin Y stain; bar – 10 μm The content of fat in goat’s milk was gradually raised while increasing of somatic cell count, but significant difference was not N. M. Zazharska 11 found. Also a positive correlation between the somatic cell count and protein and lactose was found, but in the milk with somatic cell count of more than 3 million/ml there was a sharp decrease. The opposite trend was observed concerning the freezing point of milk. The freezing temperature increased while reducing of protein in goat milk. The study of Hungarian scientists also showed a positive correlation (P < 0.01) between the somatic cell count and the protein content (r = 0.67; P < 0.001) and a negative correlation between the somatic cell count and the lactose content (r = –0.41, P < 0.001) and the freezing point (r = –0.33) in goat’s milk (Pajor et al., 2013).

Figure 4. Disadvantages in goat’s milk smears: Romanovsky – Giemza stain; bar – 10 μm; a – cytoplasmic particle, b –“shadow” of the cell, c – piece of paint There was no correlation between the somatic cell count and the content of protein, fat and lactose in goat milk according to the results of other scientists (Bagnicka et al., 2016). Contrary, polish scientists point out that there is an effect of the somatic cell count on lactose content in goat milk (Czopowicz et al., 2013).

Monitoring studies of goat’s and cow’s milk in France and Ukraine

The safety and quality of raw milk in Ukraine remained the biggest problem in retooling of dairy enterprises by newest processing lines, the introduction of modern quality control systems. In France 12 Chapter 1. Assessment of safety and quality of goat’s milk every farmer is interested in improving of product quality because it affects the price of milk. Monitoring of goat milk indexes was conducted in Ukraine, compared with similar ones in the milk laboratory of LILCO (Laboratoire Interprofessionnel Laitiere du Cen- tre Ouest – Interprofessional milk laboratory of center and west), Surgères, France. LILCO – one of 16 laboratories for control of milk quality in France. LILCO serves more than 5 thousands farmers who get milk from cows and goats. Laboratory analyses the milk from the herd of each farmer three times a month. Based on the results of milk analysis laboratory forms the price, that the dairy company has to pay to the farmer. In Ukraine, unfortunately, dairy plants do not accept goat milk for processing, although it is a valuable raw material than cow milk (Zazharska, 2015). Laboratory LILCO determines fat, protein, somatic cells count, freezing point, lipolysis, microbial contamination, inhibitors, bu- tyric bacteria in the milk of cows and goats. All physical and chemical indexes of milk were determined by devices FossomaticTM FC and MilkoScanTM FT+. To 16 thousand samples of milk per day are analyzed in the laboratory. Many methods of verification of the accuracy of analyzes, including repeatability and reproducibility, tracage, reference methods take place to control the operation of very expensive and modern apparatuses. Reference methods for fat (acid Gerber method), protein (with amide black), freezing point (by cryoscope) perform daily to control the accuracy of machines MilkoScan. There is also an internal control sample and “gamma” (10 samples with known parameters) from the reference-laboratory Ceca Lait. Microbial contamination of milk is determined by epi- fluorescence microscopy (FOSS Integrated Milk Testung BactoScan- FC), reference method – passaging through the nutrient medium (Zazharska, 2016b). Laboratory LILCO was accredited to ISO 17025 by Committee of Accreditation COFRAC (Comité français d’accréditation). Assur- ance of quality of testing and research methods of analyses was pro- vided by CNIEL (Centre national interprofessionnel de l’economie N. M. Zazharska 13 laitière – National interprofessional center of dairying). Laboratory data of 2013, 2014 years were statistically processed during the in- ternship. Laboratory analyzed per year more than 3.5 million of milk samples for determination of fat, protein, somatic cells count. Number of samples of goat milk was about 4 times less than of cow milk, the majority of samples were from individual animals. The objective of the study was to compare the parameters of cow’s and goat’s milk on the basis of analysis in French laboratory during two years. Fat content in cow milk in France was higher than in goats. In cow milk indicators of fat were almost constant throughout the year, from 4.0% in summer to 4.3% in winter (fig. 5).

Figure 5. Fat content in milk of cows and goats for two years (%, n ~ 292 thousand samples per month, according to the data of LILCO) Fluctuations of this indicator in goat milk were more significant – from 3.3% in summer to 4.5% in winter. A similar tendency was observed in regard to protein (Fig. 6). 14 Chapter 1. Assessment of safety and quality of goat’s milk

In cow milk protein content was almost at the same level (3.3%) throughout the year, decreasing slightly to 3.2% in June and July. The protein content in goat milk fluctuated greatly: from 3.7% in Janu- ary the figure gradually decreased to 3.1% in June and July, and then increased to 3.9% in December.

Figure 6. Protein content in milk of cows and goats for two years (%, n ~ 292 thousand samples per month, according to the data of LILCO) So, according to the processing of statistical data obtained in the LILCO in cow milk indicators of fat and protein were almost constant throughout the year, in goat milk – are gradually decreased in the summer. Somatic cell count in cow milk during the year was in average 307 ± 5 × 103 cells/ml; in goat milk in March – the lowest level of about 1 577 ± 77 × 103 cells/ml, and in December was 2 times more (flow cytometry method) – (Fig. 7). Infectious and non-infectious factors are influenced greatly on the somatic cells count in goat milk, unlike in cow milk. It was established that somatic cell count did not correlate with the index of bacterial contamination in goat milk. N. M. Zazharska 15

Figure 7. Somatic cell count in milk of cows and goats for two years (× 103 cells/ml, n ~ 293 thousand samples per month, according to the data of LILCO) In France, the best goat milk is considered with somatic cell count ≤ 1 million/ml. But dairy plants can take for processing goat milk with somatic cells count up to 3 million/ml at a reduced price. Ac- cording to Ukraine requirements it is permitted to 400 × 103 cells/ml for the highest quality cow milk, and for the goat milk – up to 500 × × 103 cells/ml. Perhaps we have to review Ukraini-an requirements for somatic cell count of goat milk. Bacterial contamination of cow milk was on average 22–24 × 103 CFU/ml during the period of two years (Fig. 8). Requirements for cow milk indexes in France are stricter than EU Directive concerning bacterial contamination and somatic cell count. The fluctuations of indexes of microbial contamination in goat’s milk were from 17 to 35 × 103 CFU/ml during two years ac-cording to the data of LILCO. The indexes of somatic cell count of milk did not always correlate with microbial contamination, espe-cially in goat’s milk. 16 Chapter 1. Assessment of safety and quality of goat’s milk

Freezing temperature of cow’s milk was about –0,52 ºC, of goat’s milk – about –0,55 ºC (Fig. 9). The level of lipolysis in cow’s milk varied within 0,55–0,71 mg equivalent/100 g fat. The lipolysis indexes in goat’s milk increased from March to June (to the 0.67 mg equivalent/100 g fat), and then insensibly decreased until September. The parameter of milk density is not determined in the laboratory LILCO because this indicator is considered non-informative. To determine the falsification of milk with water, they use a freezing point. The acidity of milk is also not determined, because all farmers have cooling tank, and milk collection is in accordance with the schedule. Comparative analysis of indicators of goat and cow milk in Ukraine: significant variations in acidity, density, somatic cell count of goat milk during the year were noted (Table 2).

Figure 8. Bacterial contamination in milk of cows and goats for two years (× 103 CFU/ml, n ~ 16 thousand samples per month, according to the data of LILCO) N. M. Zazharska 17

Indicators of the acidity in goat milk vary from 14 °T to 27 °T, of the density – from 25.6 °A to 35.4 °A. In milk of cows the average index of acidity is 17.6 ± 1.7 °Т, of density – 28.6 ± ± 1.3 °А. The smallest somatic cell count in the goat’s milk was observed in the autumn 265 ± 41 × 103 cells/ml, the highest – in the winter – 451 ± 46 × 103 cells/ml (Р < 0.05) (Table 2). These indexes corre-spond to the "higher" grade of goat's milk in accordance with "Goat's milk. Raw. Specifications: DSTU7006: 2009" – up to 500 × 103 cells/cm3.

Figure 9. Freezing temperature in milk of cows and goats for two years (оС, n~18 thousand samples per month, according to the data of LILCO) In cow milk the biggest somatic cell count was observed in winter 205 ± 67 ×103 cells/ml and in summer – six times less (Р < 0.05, tab. 3). These values of somatic cells correspond to the "extra" grade of cow's milk – up to 400 × 103 cells/cm3 (Zazharska, 2014; Zazhar- ska & Pryadka, 2015). 18 Chapter 1. Assessment of safety and quality of goat’s milk

Table 2. – Results of seasonal study of goat’s milk in Ukraine (x ± SE) Somatic cell count Season, num- Acidity, °Т Density, °А (by viscosimetric ber of samples method), × 103 cells/ml Spring, n = 7 17.3 ± 2.7 29.7 ± 1.5 336 ± 77ab Summer, 17.6 ± 2.8 29.2 ± 1.2 388 ± 50ab n = 12 Autumn, 21.8 ± 3.2 29.8 ± 1.9 265 ± 41a n = 53 Winter, n = 6 21.3 ± 1.6 30.5 ± 1.0 451 ± 46b Note: different letters within the column correspond to the selections which had significant differences between one another according to the results of Tukey’s test (P < 0.05) with Bonferroni correction

Table 3. – Results of seasonal study of cow’s milk in Ukraine (x ± SE) Somatic cell count Season, num- Acidity, °Т Density, °А (by viscosimetric ber of samples method), × 103 cells/ml Spring, 17.2 ± 1.0 28.7 ± 1.1 111.7 ± 29.5ab n = 15 Summer, 15.1 ± 1.3 27.9 ± 1.1 34.1 ± 8.8a n = 14 Autumn, 16.8 ± 1.2 28.4 ± 1.4 164.8 ± 17.5ab n = 12 Winter, n = 8 18.2 ± 1.3 28.9 ± 1.6 204.9 ± 67.5b Note: different letters within the column correspond to the selections which had significant differences between one another according to the results of Tukey’s test (P < 0.05) with Bonferroni correction The objective of the study was to estimate the effect of the number of lactation on the indexes of milk of cows and goats by breeds. The material of research were milk samples from 10 cows of Holstein N. M. Zazharska 19 breed and 17 goats of German white, Anglo-Nubian and Alpine breeds. The samples were analyzed on the somatic cell count, the content of fat, protein during the period of four lactations. The biochemical indexes of milk were determined by means of “Ekomilk” and “Dairy Spec Bentley Instruments”. The somatic cell count in milk samples was determined by flow cytometr “SomaCount Bent- ley Instruments”. It was established that the indexes of milk such us fat and protein depend on the quality of feeding basically, and not on the quantity of lactations. The indexes of somatic cell count in milk of cows of the 2nd, 3rd, 4th lactation was more at 5,2, 4,6 and 4,9 times respectively, according to the parameter of the first lactation (Р < 0,05). The smallest indexes of somatic cell count throughout the life of the animal was observed in the milk of the first lactation (in cows – 37 ± 7, in goats – 712 ± 174 × 103 cells/cm3, fig. 10). In cows from the second to the fourth lactation, somatic cell count was recorded at the level of 169–192 × 103 cells/cm3, in goats – 880–1092 × × 103 cells/cm3 (flow cytometry method).

Figure 10. The somatic cell count in milk of cows (n = 10) and of goats (n = 17) depending on the number of lactation 20 Chapter 1. Assessment of safety and quality of goat’s milk

In general, we did not find interconnection between increasing of the somatic cell count and the age of cows or goats. The index of somatic cell count in second lactation increased in three times compared with the first in Alpine breed goats, while in the German white and Anglo-Nubian breeds it had decreased by 5.1 and 31.7% respectively, which proves the great variability of this indicator (Zazharska et al., 2017 a).

Effect of exogenous and endogenous factors on the quality and safety of goat milk Bacterial contamination of milk at different temperatures and shelf life The main problem of the Ukrainian dairy industry is adapta- tion to European standards in quality parameters of milk. Milk in Ukraine has big bacterial contamination (second grade of DSTU – to 3 million CFU/cm3). The aim of the study was to determine bacterial contamination and physical-chemical parameters in goat milk depending of different temperatures and periods of storage. Analyses were conducted in the laboratory LILCO (Laboratoire In- terprofessionnel Laitiere du Centre Ouest – Interprofessional milk laboratory of center and west), Surgères, France. All biochemical indexes of milk were determined by devices FossomaticTM FC and MilkoScanTM FT+. Bacterial contamination of milk was determined by epifluores- cence microscopy (FOSS Integrated Milk Testung BactoScanFC). The data were analysed in Statisica 6.0 (StatSoft Inc., USA). The data in the tables are presented as x ± SE (x ± standard error). The differences between the values in groups were determined using Tukey test, where the differences were considered significant at P < 0.05 (with taking into account the Bonferroni correction). For the first experiment, 2 liters of bulk tank goat milk was selected from the farmer – the client of laboratory LILCO. Milk was divided in 45 flacons: the first 15 samples were cooled immediately and stored at 4 °C. Second 15 samples were stored at 8 °C, the last 15 flacons – at 12 °C. 5 samples of milk from every refrigerator were N. M. Zazharska 21 analyzed at 18 hours after milking, at the second and third day. The enterprises do not collect milk from farms every day, so the samples were kept for several days for research. All indexes of milk were at the same level at different temperature of storage after 18 hours of milking (Table 4). The length of bactericidal phase depends on temperature of milk storage. Bactericidal properties were in milk during the first day, so bacterial contamination was at one level, even at 12 °C. Table 4. – Results of the analysis of bulk tank goat’s milk samples, (x ± SE, n = 5) Bacterial contamination at different temperature storage of milk samples, Storage time ×103 CFU/cm3 4 °C 8 °C 12 °C First day (18 hours 23.2 ± 0.4 23.2 ± 0.8 22.6 ± 0.8 after milking) Second day 20.6 ± 0.7a 20.4 ± 0.5a 151.6 ± 3.4b Third day 21.8 ± 0.4a 28.4 ± 0.5b 1236.0 ± 55.6c Note: different letters within the line correspond to the selections which had significant differences between one another according to the results of Tukey’s test (P < 0.05) with Bonferroni correction All indexes of milk have not changed at the second day of storage compared to the first day besides a total plate count of milk which was kept at a temperature of 12 °C. Bactericidal phase finished and microbes began to multiply in milk. The bacterial contamination of milk kept at a temperature 12 °C was higher 7.4 times compared with milk stored at temperatures 4 оС and 8 оС at the second day after milking (P < 0.001). A total plate count of goat milk stored at temperature 8 °C (requirement for the delivery of milk to enterprises) was much higher than when stored at temperature 4 оС (21.8 ± 0.4 × 103 CFU/cm3) at the third day after milking (P < 0.05). If the milk has not been collected within 2 hours after milking, it should be cooled to a temperature of 8 °C or lower, or 6 °C 22 Chapter 1. Assessment of safety and quality of goat’s milk and lower if the collecting lasts more than a day (Regulation (EC) № 853/2004). It was established that bacterial contamination in goat milk stored at temperature 8 °C was much higher than in milk stored at temperature 4оС at the third day after milking (P < 0.001). So, it is not enough to cool the milk to 8 °C after milking: at third day bacterial contamination of milk is too much for producing dairy products. The maximal continuation of bactericidal phase is possible only with the rapid cooling of milk after milking to 4 °C. Ev- ery farmer in Ukraine has to milk animals only mechanically with closed supply milk to the cooling tank. Then bacterial contamination of milk will correspond to European requirements. It was proved that to ensure high quality milk it should be the rapid cooling of milk after milking to 4 °C. For the second experiment, 24 samples of cooled bulk tank goat milk were selected be transported within 2–3 hours at different temperatures. Then all samples were stored day at 4 °C. The indicators of bacterial contamination, fat, protein, freezing point, somatic cell count, urea were similar for different temperatures of transporting milk samples. It was noted the big somatic cell count (> 2000 × 103 cells/ml) at low bacterial contamination (19.6 × 103 CFU/ml) in goat milk. It was proved that milk samples can be transport to the laboratory at a temperature of 2, 10 or 20 °C during 2–3 hours if the milk after milking was cooled immediately and stored at 4 °C. For the third experiment, 5 samples of non-cooled cow’s milk were analyzed at 3 hours after milking, 5 samples of cooled cow’s milk – after a day. A total plate count of milk which was cooled and stored one day at 4 °C was in 4.6 times less (P < 0.01) than non- cooled milk, which has been analyzed in 3 hours after milking. This proves that bacterial contamination of milk in Ukraine accordance with European requirements (up to 100 ×103 CFU/ml) is possible only when rapid cooling to 4оС and storing in the cooling tank (Zazharska, 2016 a). N. M. Zazharska 23 Comparative characteristic of milk quality of German White, Alpine and Anglo-Nubian breeds of goats The comparative characteristic of milk indexes of different breeds of goat was conducted. The material for research was individual milk samples from 21 goats of Alpine, German White and Anglo-Nubian breeds. The best milk quality indexes were observed in the Anglo- Nubian breed (fig. 11) – the highest fat content (3.81%), protein (3.52%), lactose (5.25%), solids (13.33%), but the goats of the Alpine and German white breeds are characterized by large milk yields. A significant increase in the fat content in milk was found in the Anglo-Nubian and Alpine goats (P < 0.05). The indexes of calcium content ranged from 94.7 to 169.8 mg/100 g, the significant difference was found between the milk of German White (the highest calcium content – 169.8 ± 28.1 mg/100 g), and Alpine breed (P < 0.05). The content of calcium in goat’s milk is on average 124 mg/100 g (Greppi et al., 2008). The somatic cell count of milk of all goat breeds corresponded to a higher grade of DSTU7006:2009.

Figure 11. Physic-chemical indicators of milk of different breeds of goat 24 Chapter 1. Assessment of safety and quality of goat’s milk

The best suitability of milk for cheesemaking was marked in Al- pine goats. The smallest somatic cell count (271 × 103 cells/cm3) was noted in German White goats. Therefore, it is expedient to breed all three breeds of goats (Zazharska & Gramma, 2016). Influence of diet on the productivity and characteristics of goat milk We assessed the relationship between the milk quality indexes of goats of different breeds and their diets. There were used in the experiment: Anglo-Nubian (7 animals), German White (7) and Alpine goats (7). We investigated the influence of two diets: a routine diet (hay and concentrates) and a diet improved by granulated alfalfa hay, mixed feed. The volume of the morning milk yield and biochemical indexes of the milk of each goat after machine milking were measured. It was established that the milk yield of the Alpine goats increased 3 times; and that of the German White goats increased more than 2.5 times when goats got improved diet (added granulated alfalfa hay and concentrate feed). Feeding improved diet resulted in significant increase in fat content of milk of all breeds from 2.06–2.62% to 3.76–5.69 (P < 0.05). A significant increase (Р < 0.05) in the protein was observed in Anglo-Nubian (from 3.02 to 3.31%) and German White goats (from 2.88 to 3.03%) when they got the improved diet. Compared with the German White and Alpine goats, the highest figures for the fat, protein and lactose were found in milk of the Anglo-Nubian goats with the routine and improved diet. With all breeds under study the freezing point and electrical conductivity of the milk decreased when they were fed the improved diet. An inverse relationship was found between the protein content and the freezing temperature in the goats’ milk: when the protein content increased, the freezing point decreased. Content of protein in milk of Saanen and Alpine goats from 27.0 to 29.2 g/kg (on average 2.8%), and fat content equal to 30.2–34.1 g/kg (on average 3.2%) (Maurer et al., 2013). According to the other data the protein content in milk of the British Saanen goat is 2.6%, of Nubian in Great Britain – 3.6%, Alpine and Saanen in France – 3.2%, and fat content of 3.5%, 4.9%, and 3.6%, respectively N. M. Zazharska 25

(Yangilar, 2013). In our studies, a significant increase of fat and protein content in goat’s milk was recorded after improvement in the diet (Zazharska et al., 2018). Physical and chemical composition of goat and sheep milk depending on the altitude of grazing The quality and safety parameters of milk were compared, depending on the altitude of grazing of goats in the valleys of Zakarpattya (251, 309, 341, 376, 394, 524, 580 and 750 m above sea level). The milk samples were studied from 5 goats at each height. It was determined, that acidity of milk ranged from 13 to 17 oT, the density of the goat – from 24.4 oA to 30.5 oA in goats, grazed in the valleys of Zakarpattya. The protein content of goat milk was – 2.90 ± 0.07 to 3.19 ± 0.15%, fat content – 2.60 ± 0.50 to 5.61 ± 0.66%, somatic cell count – up to 359 ± 226 ×103 cells/cm3. At the highest altitude of the grazing – 750 m above sea level, a lowest fat content in the milk of goats (2.6 ± 0.5%) was marked that in 1.4–2.2 times was less than the values of other altitude of the grazing. The lowest concentration of somatic cells in goat milk was 82 ± 29 × 103 cells/ cm3 (by viscosimetric analyzer) at this altitude, that in 1.9–4.4 times was less than the values of other heights (Fig. 12).

Figure 12. Somatic cell count in goat milk depending on the altitude of grazing (by viscosimetric method) 26 Chapter 1. Assessment of safety and quality of goat’s milk

It was proved that if the fat of goat’s milk was more, then its density was less. An inverse relationship was found between the protein content in the goats’ milk and the freezing point: with increase in the protein content, the freezing point was reduced. It was revealed that somatic cell count in goat milk (359 × 103 cells/ml) was much less than requirements in Europe (< 1000 × 103/ml). The high fat content in the milk of goats (5.61%) was observed at an altitude of 341 m above sea level (P < 0.01). The highest freezing point of milk, and at the same time – low figures of protein and density were marked in the same animals. At the highest altitude of grazing – 750 m above sea level, a lowest fat content (2.6%) was marked along with highest figures of dry non-fat milk solid and protein content in goat’s milk. The lowest somatic cell count in milk of goats at this altitude indicates a high sanitary quality of milk (Fotina & Zazharska, 2016). Study of indicators of safety of goat’s milk for intense man-caused pollution The hazard of grazing dairy goats near highways and racing them along the roadside of the highways to the pasture has been proved. Content of plum (0,23 ± 0,08 mg/kg) in milk from the goats of the city of Dnipro exceeds a maximally possible level in accordance with Ukrainian requirements more than in 2 times, and maximally possible level for the milk used for production of child’s and dietary products more, than in 4 times. The content of plum and cadmium in a cottage cheese in 2.1–2.4 times is higher, and in whey in 1.5–1.6 times below as compared to milk (Fig. 13). The purpose of the study was to estimate changes of goat’s milk depending on the season and lactation period. Monitoring study of the milk from 4 goats of 1–2 lactation and 4 goats of 4–5 lactation from the village Mar’yanske, Apostol district of Dnepropetrovsk region were conducted (total 211 samples). The quality and safety indexes of goat’s milk depending on seasons were determined by analyzer of milk “Ekomilk”. The somatic cell count depending on lactation period, portions of milk during milking was determined by viscosimetric analyzer “Somatos”. N. M. Zazharska 27

Figure 13. The content of plum in goat milk and dairy products, mg/kg Effect of lactation period, yield time, season on goat milk indexes Cheese, dangerous for the content of salts of plum and cadmium is produced from such milk in milk processing enterprises. It is proved, that protein and lactose in goat milk in autumn increased on 17.5% and 13.5% accordingly compared to the summer (Р < 0.05). During summer fat (3.70 ± 0.18%) in milk was lower in 1.8 times compared with the winter period (Р < 0.05), in 1.5 times – with spring (Р < 0.05), on 16% – with autumn period (Р < 0.05). The lowest freezing point marked in winter, accompanied by a high content of lactose, fat and the largest somatic cell count. In autumn and winter the somatic cell count was 3.7 (Р < 0.05) and 5 (Р < 0.05) times accordingly more than the spring-summer figure (96 ± 14 ×103 cells/cm3). In the first month of lactation goat milk contained very low somatic cells count from 33 to 107 × 103 cells/cm3. Low figures of somatic cell count (15 to 63 × 103 cells/cm3) was marked in milk of goats of first lactation, but it lasted only seven months. The somatic cell count in milk of goats of 1–2 lactation is slightly lower than in animals of 4–5 lactation, but no statistical difference was detected (Fig. 14). During the year, the indexes of milk yield increased in May and September, decreased in summer months – because of the heat, and to the end of lactation – in 7–9 months. 28 Chapter 1. Assessment of safety and quality of goat’s milk

While studying the milk of eight goats for more than one and a half years, each animal noted an increase in the somatic cell count in the evening and in the morning. The interconnection of the index of somatic cell count and yield time was not found. The index can change twice and more at morning and evening milking during the day (Zazharska & Kostyuchenko, 2015; Shapovalov et al., 2015; Fotina et al., 2018).

Figure 14. Somatic cell count in goat milk depending on lactation month, × 103 cells/cm3 Colostrums contain large concentration of immunoglobulin G, which is particularly important in the first hours and days of a new- born. It is very important to be sure that there are no colostrums in the total yield because of the high acidity of colostrums. That milk cannot be pasteurized or sterilized. The changes of organoleptic, physical-chemical parameters and concentration of immunoglobulin G were studied in 44 samples of goat milk and colostrum depending on the period of lactation. The concentration of immunoglobulin G was analyzed by means of IDRing Plate-Caprine IgG Test, France (simple radial immunodiffusion). N. M. Zazharska 29

The organoleptic indicators (colour, consistency, taste, smell) of goat’s colostrums at first day after lambing were significantly different from milk of other days. Colostrums colour in the first 3 days was creamy-yellow, texture was viscous, especially at the first day. The maximum acidity 56 °T was in the first colostrums yield, due to the maximum concentration of immunoglobulin, at the second and third day acidity was 15–16 °T. All main parameters of colostrums significantly decreased at the second day of lactation. Fat and lactose of colostrums at the second day after lambing fell more than 2 times, dry non-fat milk solids – by 42.6%, density – 40% of total protein – 42%, the freezing point – 44% compared to first day. Conductivity and pH on the contrary increased at the second day of lactation in 44.9 and 5.7% accordingly, but these were only parameters where significant difference compared to the first colostrums yield wasn’t found for seven days. Determined, that fat, dry non-fat milk solids, density, protein, lactose of colostrums during the period from second to seventh day of lactation were significantly below from the same parameters of first milk yield (P < 0.05), which proved the value of the first colostrums Somatic cell count in colostrums on the second day of lactation decreased twice, and on the seventh – in ten times compared to the first milk yield (680±217 × 103 cells/cm3) (P < 0.05). According to data of Spanish scientists, Ig G content in goat colostrums was 19.97 g/l. The average concentration of Ig G after the seventh milking was less than 2 g/l, and after the eleventh – less than 1 g/l (Fernandez et al., 2006). The same pattern was observed in our study. The Ig G content in goat’s colostrums in the first yield was 15.79 g/l, the day after lambing – 16.8% less. The sharp decline of Ig G concentration compared with the first milk yield was from the 3rd day of colostric period – in 6 times less (P <0.05). From the 6th day of lactation the concentration of immunoglobulin G in milk wasn’t exceed 1 g/l, which indicates the possibility of adding such milk to the total yield for further processing. The system for obtain- ing high quality goat’s milk has been developed. System is due to the 30 Chapter 1. Assessment of safety and quality of goat’s milk perfect cleaning of milking equipment, proper milking and care of udder health in goats. Also, in order to improve the hygienic state of the udder and to reduce the amount of somatic cells in milk, it is recommended to use mean for pre-milking washing “MolSan” and the means for udder care: ointment “Fitosept”, “Dbayliva doyarochka”, “Zorka” cream, gel of “Nizhnodiy” (Fotina et al., 2015; Fotina & Zazharska, 2015; Za- zharska & Ryaba, 2016). N. M. Zazharska 31

References 1. Bagnicka E., Łukaszewicz M., & Ådnøy T. (2016). Genetic parameters of somatic cell score and lactose content in goat’s milk. Journal of Animal and Feed Sciences, 25(3), 210–215. 2. Barth K. (2009). Eutergesundheitsüberwachung bei Milch- schafen und Milchziegen – welche Methoden sind geeignet. Landbauforsch SH, 332, 89–95. 3. Czopowicz M., Strzalkowska N., Rzewuska M., Winnicka A., Joz- wik A., Kosciuczuk E., Kaba J., Jarczak J., Krzyzewski J., Bagnic- ka E. (2013). Factors influencing technological properties of goat milk. Goat Milk Quality Regional IGA Conference, Norway. Ab- stractgbs. 14. 4. Fernandez A., Ramos J. J., Loste A., Ferrer L. M., Figueras L., Verde M. T., Marca M. C. (2006). Influence of colostrum treated by heat on іmmunity function in goat kids. Comparative Immu- nology, Microbiology & Infectious Diseases, 29, 353–364. 5. Fotina T. I., Fotina H. A., Ladyka V. I., Ladyka L. M., Zazhar- ska N. M. (2018). Monitoring research of somatic cells count in goat milk in the eastern region of Ukraine. Journal of the Hel- lenic Veterinary Medical Society, 69(3), 1101–1108. 6. Fotina T. I., Zazharska N. M., Kostyuchenko V. Y. (2015). Influ- ence of facilities for milking on sanitary quality of goat’s milk. Visnik Sumskogo nacionalnogo agrarnogo universitetu, 37, 59–65. 7. Fotina T., & Zazharska N. (2016). Physical and chemical composition of goat and sheep milk depending on the altitude of grazing. The Animal Biology, 18(4), 106–112. 8. Fotina T. I., & Zazharska N. M. (2015). Influence of drugs for udder hygiene on sanitary parameters of goat milk // Materials of the V International Congress of Veterinary Pharmacology and Toxicology, Vitebsk, Belorus, 380–384. 9. Greppi G. F., Roncada P., Fortin R. (2008). Protein components of goat’s milk. In: Pulina G.; Cannas A. (Eds.) Dairy goats feeding and nutrition. 2.ed. Bologna: CAB International, 71–94. 32 Chapter 1. Assessment of safety and quality of goat’s milk

10. Maurer J., Berger T., Amrein R., Schaeren W., Stierli M. (2013). Critères de qualité pour le lait de chèvre et de brebis: exigences et valeurs indicatives ainsi que propositions pour un paiement du lait selon des caractéristiques qualitatives. ALP Forum, 97, 1–16. 11. Pajor F., Sramek А., Tоth G., Pоti P. (2013). Effect of somatic cell count on some chemical, physical and bacterial properties of milk in a Hungarian Alpine goat farm. Goat Milk Quality Re- gional IGA Conference, Norway. Abstracts. 38. 12. Park Y. W. (1991). Interrelationships between somatic cell counts, electrical conductivity, bacteria counts, percent fat and protein in goat milk. Small Ruminant Research, 5(4), 367–375. 13. Schüppel H. & Schwope M. (1999). Zum Gehalt somatischer Zellen und zur mikrobiologischen Beschaffenheit der Milch von Ziegen mit klinisch unauffälligem Euterbefund. Milchwis- sensch, 54, 13–16. 14. Shapovalov S., Fotina T., Kalachnikov V. and Zazharska N. (2015) Composition physico-chimique du lait de chèvre de l’Est de l’Ukraine. Revue Écologie-Environnement, Algérie 11: 70–73. 15. Yangilar F. (2013). As a potentially functional food: Goats’ milk and products. J. Food Nutr. Res., 1(4), 68–81. 16. Zazharska N. & Kostyuchenko К. (2015). Influence of lactation period, yield time, season on the somatic cells count in goat milk. Prob- lems of zooengineering and veterinary medicine, 31(2), 179–184. 17. Zazharska N. (2015). Organization of work and analysis at the milk laboratory in France. Materials of the international scientific-practical conference, Ganja Azerbaijan, 1, 480–484. 18. Zazharska N. M. (2014). Somatic cell count of cow and goat milk. News of Sumy National Agrarian University, 1 (34), 89–92. 19. Zazharska N. M. (2016a). Bacterial contamination of milk at different temperatures and shelf life. Scientific Messenger of LNU of Veterinary Medicine and Biotechnology, 70, 108–112. 20. Zazharska N. M. (2016 b). Comparative characteristics of cow`s and goat`s milk, according to the data of laboratory LILCO. Sci- entific Bulletin of the National University of Life and Environ- mental Sciences of Ukraine, 237, 297–308. N. M. Zazharska 33

21. Zazharska N. M. (2018). Сriteria for safety and quality assessment of goat’s milk. The thesis for the scientific degree of doctor of the veterinary science by specialty 16.00.09 – veterinary and sanitary expertise. Sumy National Agrarian University. 22. Zazharska N. M., & Gramma V. O. (2016). Comparative characteristic of milk quality of German white, Alpine and Anglo-Nu- bian breeds of goats]. News of Zhytomyr National Agroecologi- cal University, 1 (53), 214–220. 23. Zazharska N. M., & Pryadka E. V. (2015). Influence of lactation period, yield time, season on the somatic cell count in cow milk. Science and Technology Bulletin of SRC for Biosafety and Envi- ronmental Control of AIC, 3(1), 107–112. 24. Zazharska N. M., & Ryaba A. A. (2016). Sanitary quality of goat milk in the application of the homeopathic preparations for milking. Scientific and technical bulletin of State Scientific Research Control Institute of Veterinary Medical Products and Fodder Additives and Institute of Animal Biology, 17 (1), 72–77. 25. Zazharska N. M., Kurban D. A., & Нolubyeva O. V. (2017 a). Con- tain of fat, protein, somatic cells in cow’s and goat’s milk depending on number of lactation. Theoretical and Applied Veterinary Medicine, 5(4), 17–24. 26. Zazharska N. M., Neverkovets N. Y., & Danyliuk V. O. (2017 b). Parameters of subclinical mastitis in goats. News of Dniprop- etrovsk State Agrarian and Economic University, 45, 77–81. 27. Zazharska N. N., & Rosenko S. O. (2018). Study on the prevalence of subclinical mastitis in goat milk. Theoretical and Applied Vet- erinary Medicine, 6(3), 50–53. 28. Zazharska N. N., & Zharko L. (2016). Comparison of methods efficiency for determination of somatic cells count in goat milk. Science and Technology Bulletin of SRC for Biosafety and Envi- ronmental Control of AIC, 4(4), 29–35. 29. Zazharska N. N., Boyko O., & Brygadyrenko V. (2018). Influence of diet on the productivity and characteristics of goat milk. In- dian Journal of Animal Research, 52(5), 711–717. 34

Chapter 2. Coleoptera pests of stored food supplies and field crops

V. I. Rusynov, V. O. Martynov, T. M. Kolombar Oles Honchar Dnipro National University

Introduction1

Coleoptera is the most numerous order of insects, containing over 400 000 species, which is almost 40% of described insects and 25% of all known animals. The number of beetles in the fauna of Ukraine is still unclear, but it is assumed that it cannot be less than 30 000 species (Bartenev et al., 1997). Most species at the imago stage have a hard chitinous exoskeleton. The elytra are highly sclerotized, the hindwings are membraned with poor venation, are used for flying and while the insect is at rest, the hindwings are folded under the elytra. The wings, very rarely the elytra, can be reduced. The mouthparts are of chewing-biting type, usually are well deve loped and are differently structured in different groups. In most cases, the antennae comprise 11 segments and are different in form and length. Out of three segments of the thorax, the most developed is the prothorax, which is flexibly connected with the mesothorax, which is flexibly connected to the metathorax. Each of these segments is divided into the upper part – the tergum, the lower part – the sternum, and the lateral – the pleura. The legs are well developed, typically they are running or curso- rial, but depending on the way of life, they are also saltatorial, fos- sorial, natatorial, raptorial, and adhesive. The legs usually consist of 3–5, sometimes 2 segments.

1 Rusynov V. I., Martynov V. O., & Kolombar T. M. (2019). Coleoptera pests of stored food supplies and field crops. In: Current problems of agrarian industry in Ukraine. Accent Graphics Communications & Publishing, Vancouver, Canada. – P. 34–133. Doi:10.15421/511903 V. I. Rusynov, V. O. Martynov, T. M. Kolombar 35

The abdomen is fixed, tightly connected to the metathorax, without appendages. The total number of abdominal segments is no higher than 10, each of which consists of two half rings – dorsal (tergum) and abdominal (sternum). The terga are covered by the elytra and are poorly sclerotized, and the uncovered terga are as thick as the sterna. The terminal (apical) segment of the open terga is called the pygidium, the terminal sternum – anal, its structure is different among the sexes. The coloration of beetles is diverse. It depends ei- ther on the presence in the cuticle of pigments (non metallic tones), or on a special microscopic structure (metallic tones) or on the combination of the pigments and the structures. The development of beetles involves full transformation. The eggs are usually oval, half-transparent, with thin non-pigmented cases. The larvae are quite different in structure (oligopod, polypod and wireworm types) with well developed, highly sclerotized heads and gnawing-chewing mouthparts. The thoracic legs are developed or reduced and can even be absent (larvae of snout beetles). Most larvae live a secretive life. Larvae of leaf beetles and representatives of some other families live openly on plants. The pupae are free, soft, with thin cases; pupae of most species develop discretely in soil, wood and other suitable places. Beetles inhabit all terrestrial biotopes, fresh water and to a lesser extent saline waters. Many of them are typical inhabitants of the upper layers of soil and dead and decomposing plants, some live in manure and the corpses of animals (Brygadyrenko, 2016). Especially numerous are the inhabitants of different parts and organs of plants (leaves, flowers, fruits, shoots, bark, wood, and roots). According to the type of feeding, the beetles are classified as predatory (both polyphagous and specialized) and herbivorous (phytophages). Among the latter, there are consumers of leaves (fo- livores), roots (rhizophages), wood and bark (xylophages), flowers (florivores), fruits (carpophages) and seeds (granivores). The consumers of fungi are the group mycophages, consumers of decomposing plant and animal remains – saprophages and necrophages. 36 Chapter 2. Coleoptera pests of stored food supplies and field crops

The regional fauna is represented by two suborders: the mainly predatory Adephaga and the omnivorous Polyphaga. Many representatives of the Polyphaga suborder, and, in some cases, Adephaga cause significant damage to agriculture and forestry (Vasilev, 1987). Coleoptera also includes pests of stored materials of plant and animal origin (Ptinidae, Dermestidae, some darkling beetles (Tene- brionidae), seed beetles (Bruchinae), wheat weevils (Sitophilus granaries), and Bostrichidae).

Suborder Adephaga Family Carabidae The family Carabidae including ground beetles, is an incredibly diverse group of insects that can play a major role in agroecosystems. By consuming a variety of insect pests and weed seeds, they can help protect crops from pest damage and associated losses, and decrease costs associated with pest control (Namaghi, 2011). Many species of ground beetles are predators. They feed on insects, their larvae, mollusks, worms and other invertebrates. Some eat both plant and animal food (omnivores), some ground beetles are herbivorous (phytophages) or feed on remains of animals and plants (saprophages) (Thiele, 1977). Phytophages include pests harmful to agriculture. Among predators, recourse to phytophagy is related to rapid decrease or disappearance of animal food, caused by different reasons. One of them is the death of invertebrates which are food for the ground beetles. Another cause is various viral, bacterial and zoopar- asitic diseases, or consumption of ground beetles’ potential victims by other animals. A no less important factor is the seasonal migration of the ground beetles’ prey, mostly insects. Ground beetles cannot always follow their migration. A certain negative impact is also caused by farming activity (cutting down forests, chemical processing of plants, etc.) and industry causing ground beetles to migrate to other places or turn to saprophagy and phytophagy. For example, in May 1968, in Ukraine (Vinnytsia Oblast), Clivina fossor beetles V. I. Rusynov, V. O. Martynov, T. M. Kolombar 37 destroyed 50% of seedlings of corn on farms where insecticides had been applied to the soil prior to planting, leading to the death of many insects. All herbivorous ground beetles distributed in the territory of Ukraine are polyphages. Accordingly, they are observed to have a preference for some particular plants. Sometimes an apparent relationship between beetles and certain groups of plants is explained by the fact that certain species of ground beetles are associated with particular habitats. For example, meadow ground beetles harm meadow grasses (Gramineae) and sedges (Cyperaceae), forest ground beetles harm sprouting seeds of trees, shrubs and herbaceous plants in forests, glades and forest edges. They eat ripening seeds, fruits, young leaves, root collar and other organs of the plants they damage. Many authors (Novák, 1967; Frank, 1967; Frank, 1971; Thiele, 1977) have mentioned the significant contribution of ground beetles to natural destruction of pests and forest insects, and that they sometimes even play a leading role in this compared to other predators (for example, Staphylinidae). Therefore, if necessary, plants should be protected against ground beetles using first of all agrotechnical and other environmentally friendly methods, using insecticides only in extreme cases if there is a threat of significant economic damage. When considering measures against ground beetles, one should take into account that most species of this family, except a few obligate phytophages, such as Zabrus tenebrioides and Z. spinipes, are notice- ably, sometimes significantly useful in destroying different pests of agricultural crops.

Subfamily Brachininae

Brachinus (Brachinus) crepitans Linnaeus, 1758. Size – 6.5–10.0 mm. Is characterized by visible number of abdominal sterna, equaling 8 for males, and 7 for females. The apices of the elytra are blunted, with no lateral margin. It has one supraorbital seta. The front legs of males are insignificantly widened. The color is bright: the head and pronotum are reddish, the elytra are blue-green, rarely 38 Chapter 2. Coleoptera pests of stored food supplies and field crops black-blue, of metal color. The pronotum has no carina (keel). The wings are well developed. The leathery margin on the apices of the elytra has long, downcurved setae. The elytra are usually of one col- or, metallic. The abdomen, metathorax and pygidium are black or dark brown, the base of the abdomen sometimes is reddish. The elytra are usually enlarged at the back. The intervals of the elytra are not raised, the leathery membrane on the dorsum is narrow, brown or black. The 3rd and 4th segments of the antennae are completely reddish. Cosmopolite, occurs throughout Ukraine (Brigic et al., 2009). Predator, but can damage generative organs of rye, wheat, timothy (Phleum pratense), and foxtail grass (Alopecúrus) (Eyre et al., 2009). Brachinus (Brachynidius) explodens Duftschmid, 1812. Small- er than 4.5–6.5 mm. The elytra have no pre-scutellum spot; their intervals not raised, the striae are not clearly distinct. The head with the eyes is wider or nearly wider than the pronotum. The abdomen and pygidium are black or brown. They occur in the south forest steppe and in the steppe. Broadly distributed European species, common throughout Ukraine (Saska and Honek, 2004). The beetles can harm generative organs of rye, wheat, timothy, and foxtail grass (Eyre et al., 2009).

Subfamily Harpalinae

Acinopus (Acinopus) laevigatus Ménétriés, 1832. Body size – 11–16 mm. The body is large, black, the antennae and the legs are brown. The head is slightly narrower than the pronotum. The margin of the elytra is nearly straight. The upper part is glossy. The incisure of the right mandible starts at the front border of the labrum and is covered by the labrum in the posterior part. The striae on the elytra are very thin. Distributed in Southern Europe, Eastern Medi- terranean, Central Asia, the Near East and in North-East Africa. In Ukraine, it occurs in the steppe zone, causes damage in Kherson and Odessa Oblasts and in the Crimea (Avgin and Özdikmen, 2008). Causes damage to grain (wheat, rye, corn, barley, panic grass, oats), technical (beet), garden (carrot), and medical (plantains (Plantago) V. I. Rusynov, V. O. Martynov, T. M. Kolombar 39 crops. It feeds on the wild-growing species of herbaceous plants (Po- aceae, Chenopodioideae, Compositae). Beetles and larvae are omnivores. These beetles cause damage to grain crops by eating sprouting seeds, generative organs and ripening seeds (Namaghi, 2011). Acinopus (Acinopus) picipes Olivier, 1795. Body size – 12–17 mm. Their distinctive feature is that the outer incisure of the right mandible starts near the posterior edge of the labrum. Has distinct striae on the elytra. Distributed in Europe, Western and Eastern Mediterranean, the Near East. In Ukraine, it occurs in Odessa, Mykolaiv, Kherson, Zaporizhia Oblasts and the Crimea (Fadda et al., 2008). Causes damage to grain (wheat, rye, barley, panic grass, corn) and technical (beet) crops, and also feeds on 6 species of wild-growing grass (Poaceae, Compositae). Anisodactylus (Pseudanisodactylus) signatus Panzer, 1796. Body size – 11–14 mm. Body is thickset. The head has a red spot on the forehead. The apical spur on the protibia is simple. The dorsal surface is black or with poor metallic gloss. The third interval of the elytra has no setiferous pore. The elytra are jet-black, often with bronze gloss, the epipleura are reddish. Distributed in Europe, Central Asia, China and Japan (Fazekas et al., 1997). Is common throughout Ukraine. The greatest damage is observed in the forest-steppe, subzone of the northern steppe and the foothills in the Crimea. Pantophagous. Causes damage to grain (wheat, rye, corn, barley, panic grass), grain legumes (peas), technical (beet, potatoes, mustard, chufa sedge (Cyperus esculentus), flax Linum( ), sunflower), garden (onion, lettuce, cucumber) and forage (clover, lucerne (Med- icago sativa), sainfoins (Onobrychis) crops, berries of strawberries (Fragaria), fallen fruit in gardens, seedlings of hornbeams, maple, and ash (Talmaciu and Talmaciu, 2011). Ditomus eremita Dejean, 1825. Body size – 7–11 mm. The body is thickset, the head is of the same width as the pronotum, which is significantly transversal. The color is black, and the antennae, tibiae and feet are brown. The head has unclear roundish forehead indentations. The intervals on the elytra have two irregular rows of punctures bearing protruding hair. Distributed in the south of the 40 Chapter 2. Coleoptera pests of stored food supplies and field crops

European part of Russia, Central Asia, and Eastern Mediterranean. In Ukraine, it inhabits the steppe, especially the southern part. Phy- tophage. Beetles consume sprouting seeds and young shoots of wild- growing grass (Chenopodioideae, Compositae), rarely wheat and corn (Nekuliseanu and Matalin, 2000). Harpalus (Harpalus) affinis Schrank, 1781. Body size is average – 9–12 mm, usually cylindrical. The side edge of the pronotum has only one setiferous pore near the middle. 2–3 external intervals of the elytra are densely punctured and bear thin hairs. Above it is of bright metal green, bronze and copper color, rarely is blue or black, the antennae and legs are yellow, sometimes the femora are jet-black. The larva has mandibles with one middle tooth. The front edge of the head between the two mandibles has 5 teeth, of which the middle one is the biggest, triangular with 6–12 denticles along the edge. The external side of the mandibles has one long and 3–4 short setae. The middle tooth on the front edge of the head bears 10–12 denticles (Bey-Bienko, 1965). Broadly distributed in Europe, occurs in the Near East, and in the Australasian region (Honek et al., 2003). The beetles feed on food of both animal and plant origin, and are mostly predatory. They cause damage to grain (wheat, corn, barley), grain legumes (peas, soybeans), technical (beet, mustard, flax), medicinal (plantains), garden (lettuce, cucumbers) and forage (timothy, lucerne) crops. Of other plants, it causes damage to strawberries, hornbeam, maple, spruce, and pine. Feeds on five species of wild-growing herbs (Eyre et al., 2013). Harpalus (Harpalus) distinguendus Duftschmid, 1812. Body size – 9–11 mm. The upper part of the 7th and 5th intervals of the elytra bears no large punctures. The shoulders are angular, with an obvious but often small shoulder tooth. The pronotum between the posterior edges and the main fossae is not flattened, the lateral margin is entirely narrow. Above it is metal-green, copper, more rarely blue or black-blue, the antennae are reddish-brown with red first section and infuscated bases of the second and the third sections, the legs are jet-black with brown feet. The larva has a rectangular middle tooth on the front edge of its head, with 6 small teeth. Distributed in Europe, V. I. Rusynov, V. O. Martynov, T. M. Kolombar 41

Mediterranean and the Near East, in the European part of Russia, Siberia and Central Asia (Urbaneja et al., 2006). Occurs throughout Ukraine, damage observed all over the country. Causes damage to grain (corn, oat, rye, wheat, panic grass, barley), grain legumes (peas, wild beans (Phaseolus)), medicinal (wallflower (Erysimum), poppy, restharrow (Ononis), tansy (Tanacetum vulgare), forage (clover, lucerne, sainfoins, vetch (Vicia) crops, strawberries, fallen fruit in gardens, pine, maple, hornbeam, ash (Pustai et al., 2015). Harpalus (Harpalus) froelichii Sturm, 1818. Body size is 8–10 mm. The distinctive features are as follows: the dorsum is protuberant, the antennae are reddish, the legs are jet-black, with reddish feet. Distributed in Europe, the European part of Russia, Si- beria, Kazakhstan (Bezděk et al., 2006). Pantophagous. The beetles sometimes cause damage to the seedlings, generative organs and ripening seeds of grain (wheat, barley, maize), technical (mustard, flax, cannabis), medicinal (daisy) crops and fallen fruit in gardens. Harpalus (Harpalus) fuscipalpis Sturm, 1818. The body size is 7–9 mm. The pronotum is narrowed towards its base, in front of the posterior edges is not sinuate. The upper side is flattened, the antennae are black, with reddish first section, the legs are dark, with reddish feet. Distributed in the middle belt of Europe, the European part of Russia, Siberia, Central Asia. Occurs throughout Ukraine (Bolgarin, 2010). Pantophagous. Sporadically causes harm to the seedlings, generative organs and ripening seeds of grain (wheat, rye), medicinal (daisy) and forage (lucerne) crops and fallen fruit in gardens. Harpalus (Harpalus) hirtipes Panzer, 1796. Body size – 12.5–15.5 mm. The outer angle of protibiae are cut slantwise (the outer angle is greater than the inner one), the external upper angle is stretched in a similar manner to a rotor blade. Width of the pronotum is over 1.5 times larger than its length; its sides are widely fringed in front, the widest part is in the middle, is slightly narrowed in near the base. The mesofemora have 12–15 setae along the posterior edge. Black, the antennae are brown-reddish near the top (Bey-Bienko, 1965). Distributed in the middle belt of Europe, the European part of Russia, Kazakhstan and the Caucasus (Bezděk et al., 2006). Oc- 42 Chapter 2. Coleoptera pests of stored food supplies and field crops curs throughout Ukraine. The beetles cause damage to the seedlings, the blooming parts and ripening seeds of grain (wheat, rye, barley), technical (cannabis) crops, and berries of strawberry. Harpalus (Harpalus) picipennis Duftschmid, 1812. Body size is 5–7 mm. The pre-scutellum stria on the base has no setiferous puncture. The third interval of the elytra has no spot after the middle. The angles of the pronotum are rounded, the antennae are reddish-yellow. The shoulder angle of the elytra is blunted, but obvious, with a notable edge. The top is clearly pebbly in texture; the main fossae of the pronotum are punctured. The metafemora have 6–7 setae on the posterior side. Distributed in Europe, the Euro- pean part of Russia, Western Siberia (Schwerk and Szyszko, 2009). Occurs throughout Ukraine. Pantophagous. The beetles sporadically harm the sprouting seeds, more rarely the flowering parts of grain (wheat, panic grass), medicinal (plantains), vegetable (carrot, garden asparagus), forage (clover, lucerne) crops and berries of strawberry. Harpalus (Harpalus) serripes Quensel in Schonherr, 1806. Larger 9–12 mm, wide and bulging. The upper side is black, rarely with bluish gloss. Distributed in Europe, in the Near East, in the European part of Russia, Siberia, Central Asia. Occurs throughout Ukraine. Pantophagous. Sporadically causes damage to the seedlings, generative organs and ripening seeds of grain (wheat, rye), medicinal (daisy), forage (lucerne) crops and fallen fruit in gardens (Taboada et al., 2004). Harpalus (Harpalus) servus Duftschmid, 1812. Body size is 8–9 mm. The femora are infuscated. The upper side is black. The base of the pronotum is cut with a slight groove, the posterior angles are almost acute-angled, and are blunted at the apices, the widest part is in the base or remotely behind the middle. The antennae, legs and often the base of the tibiae are red. Distributed in Europe, the Mediterranean, Central Asia, the European part of Russia and Western Siberia (Kataev, 2015). In Ukraine, it can be found in the steppe and forest-steppe. Pan- tophage. The beetles sometimes cause damage to the seedlings, generative organs and ripening seeds of wheat, barley and panic grass. V. I. Rusynov, V. O. Martynov, T. M. Kolombar 43

Harpalus (Harpalus) smaragdinus Duftschmid, 1812. Body size is 9–11 mm. The pronotum between the posterior angles and the main fossae is flattened or slightly indented, forming a significant widening of the lateral margin. Jet-black, the margins of the pronotum are reddish in the light, the elytra of males are glossy, green or blue, the elytra of females are mat, jet-black, the antennae and legs are reddish-yellow. In larvae have the front edge of the head is set between the mandibles with two pairs of lateral teeth and a rounded serrated projection between them. Distributed in Europe, the European part of Russia, Western Siberia, Central Asia (Desend- er and Bosmans, 1998). Can be found throughout Ukraine, damage observed in Polesia and in the forest-steppe. Causes damage to grain crops, grain legumes, strawberry, fallen fruit, sprouts of pine, hornbeam, maple, and ash. Harpalus tenebrosus Dejean, 1829. Body size is 9–10 mm. The third interval of the elytra has only one setiferous pore behind the middle, which is near the second stria. The pronotum is slightly narrower than the elytra, flattened, its base is slightly and not densely punctured. The episterna of the metathorax are twice as long as their width. Black, the upper side has a blue and violet gloss, the antennae are red, and their second and third sections are often infuscated, the legs are red-brown. Distributed in Central Europe, the Mediterra- nean, the European part of Russia, Central Asia (Kataev, 2013). In Ukraine, can be found in the steppe zone. Pantophagous. The beetles sometimes cause damage to sprouting seeds of maize, wheat, rye and berries of strawberry. Harpalus (Harpalus) zabroides Dejean, 1829. Body size – 12–14 mm. The antennae and legs are at least partly brown-black. The protibiae are cut straight on the top. The width of the pronotum is slightly bigger than its length, the sides have narrow margin in the front. Black, glossy, the elytra of females are mat, the antennae are red with dark bases of 2–4 sections. The larvae have a protuberant front edge of the head between the mandibles in the middle part. Distrib- uted in Europe, Western and Eastern Mediterranean, middle belt of the European part of Russia, Siberia to Transbaikalia, Austria, Belarus, 44 Chapter 2. Coleoptera pests of stored food supplies and field crops

Bosnia and Herzegovina, Bulgaria, the Czech Republic, France, Ger- many, Hungary, Italy, Moldova, the Near East, Poland, Romania, Slo- vakia, Spain, Switzerland, former Yugoslavia. Can be found throughout Ukraine, but more often in the steppe zone. (Khobrakova, 2008). In Ukraine, three zones of damage are distinguished: heightened – Odessa, Mykolaiv, Kherson, Dnipropetrovsk, Zaporizhia, and Do- netsk Oblasts, unstable – Kirovograd, Poltava, and Kharkiv Oblasts, insignificant – Northern and Western Forest-Steppe. It damages grain (wheat, rye, panic grass, oat, corn, barley, buckwheat), technical (beet, castor bean (Ricinus communis), sunflower) and forage (clover, lucerne) crops, strawberry, and fallen fruit in gardens. Microderes (Microderes) brachypus Steven, 1809. Body size is 7–10 mm. The head is notably thickened, with long setae in the front. The mentum has no middle tooth. The body is jet-black, the antennae and legs are rust-red. The bases of the pronotum are punctured, the main impressions are flattened. Distributed in the Near East, the North Caucasus and Central Asia (Andersen, 2005). In Ukraine can be found in Odessa, Kherson, Zaporizhia, Donetsk Oblasts and the Crimea. In Donetsk Oblast, the beetles have been observed to eat ripening seeds of wheat. Ophonus (Hesperophonus) azureus Fabricius, 1775. Body size – 6–8 mm. The pronotum, elytra and the upper part of the legs are densely punctured and covered with hairs. The posterior angles of the pronotum are smoothened or rectangular; if rectangular, the striae of the elytra are not punctured. The upper side or at least the elytra are metal blue or green. The legs are completely red-yellow. The pronotum is coarsely and not densely punctured. Distributed in Europe, Western and Eastern Mediterranean, North Africa, and the Near East, Central Asia, the middle belt of the European part of Russia. Can be found throughout Ukraine. Pantophagous. The beetles sporadically damage the sprouts and generative organs of grain (wheat, oat), technical (mustard), garden (carrot, sorrel, lettuce), medicinal (St John’s wort (Hypericum perforatum), plantains) crops; feeds on 7 species of wild-growing herbs (Poaceae, Composi- tae) (Honek and Jarosik, 2000). V. I. Rusynov, V. O. Martynov, T. M. Kolombar 45

Ophonus (Ophonus) sabulicola Panzer, 1796. Body size – 13–17 mm. The hairs on the elytra are protruding. The head is not flattened. Lateral margins of the pronotum are rounded. The elytra are usually of metal color. The pronotum is ordinarily significantly narrowed towards the rear, the posterior angles are blunted, but obvious. The elytra are blue or blue-green, the antennae are red-yellow, the rest of the body is black or jet-brown (Giglio et al., 2003). Dis- tributed in Europe, Western and Eastern Mediterranean, the Near East and the south of the European part of Russia. In Ukraine, it can be found in the steppe zone, including the plains in the Crimea, where it has been observed to damage sprouting seeds of wheat and panic grass. Ophonus calceatus Duftschmid, 1812. Body size – 12–15 mm. The elytra are sparsely punctured and covered with hairs. Black or jet-black, glossy, the antennae and legs are red-brown. Distributed in Europe, Mediterranean, Central Asia, the European part of Rus- sia, Siberia (Popovic, 2014). In Ukraine, can be found throughout the territory, having three zones of damage: heightened – Odessa, Mykolaiv, Kherson Oblasts and the Crimea; unstable – Kirovohrad, Dnipropetrovsk, Zaporizhia and Donetsk Oblasts; insignificant damage – forest-steppe and Polesia. Damages grain (wheat, rye, panic grass, barley, sorghum, oat, rice, corn), technical (flax) and forage (green foxtail (Setaria viridis), foxtail millet (Setaria italica) and others) crops, fruits of strawberries, fallen fruit in gardens; feeds on 10 species of wild-growing grasses, maple seeds. Ophonus griseus Panzer, 1796. Body size – 9–12 mm. The posterior angles of the pronotum are rectangular, blunted at the top. The abdomen is covered with small punctures in the middle, is smooth on the sides. Distributed in Europe, Mediterranean, Cen- tral Asia, Japan, the European part of Russia, Siberia. Can be found throughout Ukraine. The damage was observed mostly in Zakarpat- tia Oblast, Polesia, and the forest-steppe, and only in irrigated lands in the steppe (Bolgarin, 2010). Damages grain (wheat, rye, barley, panic grass, oats, corn), grain legumes (pea, wild bean), technical (beet, mustard, flax, cannabis), garden (carrots, onions, cucumbers, 46 Chapter 2. Coleoptera pests of stored food supplies and field crops lettuce), medicinal (wallflower, poppy, plantain, valerian) and forage (vica, lupin (Lupinus), lucerne) crops, and other plants such as strawberry, hornbeam, and maple (Kocourek, 2013). Feeds on 10 species of wild-growing herbs (Poaceae, Chenopodioidae, Umbel- liferae, Compositae). Ophonus hospes Sturm, 1818. Body size – 11–14 mm. The front angles of the clypeus on each side have only one setiferous pore. The elytra of females are mat, entirely dense and finely punctured, often glossy in males. The pronotum is punctured only in the posterior angles and along the lateral margins; the head is smooth. The elytra of females are entirely covered with dense punctures, whereas the elytra of males often bear punctures only on the sides. The upper side is usually blue or green, more rarely jet-brown with blue gloss, the legs are jet-black or reddish (Bey-Bienko, 1965). Distributed in Europe, Western and Eastern Mediterranean, the European part of Russia. In Ukraine, can be found mostly in the steppe and sporadically in the western forest-steppe. Pantophagous. The beetles sometimes damage the seedlings of wheat, foxtail (Setaria), tubers of chufa, young shoots of potatoes (Gailis, 2018). Ophonus stictus Stephens, 1828. Body size – 11.5–15.0 mm. The pronotum is narrowed to the rear, its posterior angles are rounded. The hairs on the elytra are dense, black. The upper part is green or blue-green, the lower part is black, the antennae and legs are yellow-reddish. Distributed in Central Europe, Western and partly the Eastern Mediterranean (Balkans), European part of Russia, Western Siberia, and in the Caucasus (Guéorguiev, 2008). In Ukraine, can be found mostly in the steppe and sporadically in the forest-steppe. Pantophagous. The beetles sometimes cause damage to the sprouts, generative organs, ripening seeds of mustard, wheat, carrots, feed on 8 species of wild-growing herbs (Poaceae, Plantaginaceae, Umbel- liferae, Compositae). Ophonus rufipes Degeer, 1774. Body size – 11–16 mm. The top of the head and the front part of the pronotum are smooth; the temples are bold or with a few hairs. The posterior angles of the pronotum are sharply rectangular or blunted at their apices. The V. I. Rusynov, V. O. Martynov, T. M. Kolombar 47 entire elytra bear dense punctures and hairs. The upper side is jet- black, the antennae and legs are reddish-yellow. The central part of the abdomen is bald and smooth, with rare punctures and hairs on the sides. Distributed in Europe, the Mediterranean, Japan, the European part of Russia, Central Asia (Matalin, 1994). Distribut- ed throughout Ukraine, the zone of constant damage includes the southern steppe, the zone of sporadic harm – the northern steppe. Causes damage to grain (wheat, rye, panic grass, barley, oat, rice, sorghum, maize, buckwheat), grain legumes (peas, wild beans (Phaseolus), soybeans, broad beans (Vicia faba), technical (beet, potatoes, sunflower, peanuts, mustard, rape (Brassica napus), chufa, castor bean, thymes (Thymus), cannabis), vegetable (tomatoes, carrots, onions, sorrel, lettuce), medicinal (wallflower, poppy, sage (Salvia), tansy (Tanacetum), plantain, restharrow (Ononis) and forage (Sudangrass (Sorghum × drummondii), timothy, vetches, lupin, clover, sainfoins, lucerne) crops, berries of strawberry, fal len fruit in gardens. Causes damage to trama ( flesh ) of edible mushrooms (penny bun (Boletus edulis), Russula), seedlings of spruce, pine, hornbeam, maple, ash and generative organs of 19 species of wild-growing herbs (Poaceae, Liliaceae, Urticaceae, Po- lygonaceae, Chenopodioideae, Leguminosae, Compositae) (Eyre et al., 2013).

Subfamily Platyninae

Amara (Amara) communis Panzer, 1797. Body size – 6–8 mm. The body is thickset, there is no puncturing on the episterna of the prothorax, and the sides of metathorax have fine punctures. The elytra of both sexes have obvious microsculpture. The upper side is bronze, rarely bronze-black. Distributed in Europe, Western and Eastern Mediterranean, the European part of Russia, Siberia and Central Asia, introduced in North America (Majka, 2005). Distrib- uted throughout Ukraine, found sporadically in the steppe zone. Pantophagous. The beetles sometimes eat sprouting seeds of wheat, flax, trauma of Russula and other edible mushrooms. 48 Chapter 2. Coleoptera pests of stored food supplies and field crops

Amara (Amara) eurynota Panzer, 1796. Body size – 9–12 mm. The pre-scutellum stria of the elytra has a setiferous pore in the front. The striae of the elytra along all its length to the top are not deep, the intervals are flat. The basis of the pronotum has no or few punctures. Above it is bronze, more rarely greenish or bronze-black; the legs are usually entirely black; three main segments of the antennae are reddish. Distributed in Europe, the Mediterranean, North Africa, the European part of Russia, Siberia and in the Near East (Saska and Honek, 2003). Distributed throughout Ukraine. Pantophagous. The beetles and larvae sporadically damage the generative organs and sprouting seeds of grain (wheat, panic grass, barley) and forage (clover, vetches, lucerne) crops, 6 species of wild-growing herbs (Poa- ceae, Urticinae, Compositae) and also young cabbage-plants and fallen fruit in gardens. Instead of strict phytophagy, the larvae prefer combined feeding on grains and other insects (Saska, 2004). Amara (Amara) famelica Zimmermann, 1832. Body size – 7.0 – 8.5 mm. The tibiae are jet-brown or black, the second segment of the antennae is infuscated on the upper part. The main impressions on the pronotum are large and deep, with practically no punctures. The seventh interval of the elytra near the apex has two punctures. The upper side is bronze or bronze-black. Distributed in Europe, Mediterranean, Northern China, North Africa, the European part of Russia, Siberia, Central Asia and the Near East (Heliola et al., 2008). Distributed throughout Ukraine. Damages edible mushrooms (Rus- sula and other), grain (wheat, rye, corn, oat, panic grass), technical (flax, potatoes), garden (lettuce, sorrel), berry (strawberry) and medicinal (flax, sage) crops, fallen fruit in gardens, tree species (hornbeam, maple, ash); feeding observed on 15 species of wild-growing herbs (Poaceae, Urticinae, Chenopodioideae, Crucíferae, Rubiaceae, Compositae). Similarly to other thermophiles, is observed in areas heated by the sun (Venn et al., 2013). Amara (Amara) familiaris Duftschmid, 1812. Body size – 5–7 mm. The pre-scutellum puncture furrow is developed normal- ly or at least there are obvious signs of it. The legs are of uniform yellow color. The upper part is dark-bronze-green, rarely blue or V. I. Rusynov, V. O. Martynov, T. M. Kolombar 49 black-bronze; 3–4 main segmemts of the antennae are red-yellow. The front edge of the pronotum is obviously sinuate, the front angles are prominent, usually sharpened. Distributed in Europe, the Mediterranean, North Mongolia, the European part of Russia and Siberia (Sotherton, 1985). Distributed throughout Ukraine. Pan- tophagous. The beetles eat generative organs of 8 species of wild- growing herbs (Poaceae, Cyperaceae, Caryophyllaceae), more rarely grain crops (wheat, rye, corn), technical (beet) and medicinal (plantain) plants. Amara (Amara) lucida Duftschmid, 1812. Body sizes – 4.5 – 6.0 mm. The front border of the pronotum is rectilinear, the front edges are blunted. The pre-scutellum stria is sometimes barely visible. Distributed in Europe, the Mediterranean, North Africa, the European part of Russia, Siberia (Tyler, 2007). Can be found throughout Ukraine. Pantophagous. The beetles eat generative organs of 8 species of wild-growing herbs (Poaceae, Cruciferrae), and less commonly grain crops (wheat, maize). Amara (Amara) ovata Fabricius, 1792. Body size – 9–11 mm. The lateral edges of the pronotum in the main part are rounded; the setiferous pore in its posterior angles is remote from the lateral edge at a distance of no less than the diameter of the pore; its base is almost or completely unpunctured. The upper part is black-bronze or bronze- green, rarer copper or blue. Distributed in Europe, Japan, the West and East Mediterranean, the European part of Russia and Siberia (Thomas et al., 2001). Can be found throughout Ukraine, but sporadically in the steppe zone. Pantophagous. The beetles and larvae sometimes eat the trama of penny bun, Russula, sprouting seeds of technical (flax, mustard, Camelina) and garden (radish (Raphanus raphanistrum subsp. sativus), turnip rape (Brassica rapa), lettuce) crops, hornbeam and young shoots of potatoes. Amara (Amara) similata Gyllenhal, 1810. Body size – 7.5–9.5 mm. The lateral borders of the pronotum in the main part are almost parallel; the setiferous pore in its posterior angles is positioned at the very lateral edge; its base is obviously punctured. The upper side is bronze or bronze-green. Distributed in Europe, the 50 Chapter 2. Coleoptera pests of stored food supplies and field crops

West and East Mediterranean, the European part of Russia, Siberia and Central Asia (Jorgensen and Toft, 1997). Distributed throughout Ukraine (Faly et al., 2017). Causes damage to grain (wheat, rye, corn, barley, flax, buckwheat), grain legumes (peas, wild beans), technical (beet, flax, mustard) garden (lettuce, cabbage, cucumber), medicinal (plantain, daisy) and forage (lupin, vetches) crops, strawberry, hornbeam, ash, maple, fallen fruit in gardens; feeding observed on 12 species of wild-growing herbs (Poaceae, Cyperaceae, Caryophyl- laceae, Cruciderae, Compositae). Amara (Bradytus) apricaria Paykull, 1790. Body size – 6.5–8.0 mm. The groove in the front of the posterior angles of the pronotum is obvious. The body is narrow and elongated. The punctures in the striae of the elytra are less notable in the back, but almost or completely absent on the upper incline. The mesothorax and the abdomen bear no punctures on the sides or bear fine punctures. The tooth of the mentum is big and large, blunted in the top. The striae of the elytra have deep punctures in the base. The elytra are 1.5-1.6 times longer than their width. The base of the pronotum has coarse but sparsely arranged punctures. The metatibiae of males have dense hairs along the internal edge. Jet-brown, often with light bronze gloss, the lower part is rusty red. Distributed in Europe, the West and East Mediterranean, Central Asia, the European part of Russia, Siberia, Central Asia (Eyre, 1994). Can be found throughout Ukraine, damage has been mainly observed in the steppe zone. Damages grain (wheat, rye, maize, barley, oat, panic grass, buckwheat), grain legumes (pea, wild bean), technical (beet, sunflower, peanut, mustard, flax, cannabis, thyme), garden (onion, carrot, sorrel), medicinal (wallflower, St John’s wort, sage), forage (vetches, clover) crops, berries of strawberry, maple, ash, fallen fruit in gardens; feeding observed on 10 species of wild-growing herbs (Poaceae, Chenopodioideae, Compositae). The beetles of this genus, similarly to Curtonotus and Harpalus genera, are known as pests of seeds of grain crops (Eyre et al., 2013). Amara (Bradytus) crenata Dejean, 1828. Body size – 6.5–8.5 mm. The punctures in the striae of the elytra are deep to the V. I. Rusynov, V. O. Martynov, T. M. Kolombar 51 top. The mesothorax and the abdomen have coarse punctures on the sides. The body is narrow. The tooth of the mentum is small, two- forked. The metatibiae of males have no dense hairs on the internal side. Brown or jet-brown, the underbody is rusty red. Distributed in the middle belt of Europe, the West and partly the East Mediter- ranean (Balkans), the middle belt of Russia and in the Caucasus. In Ukraine, can be found mostly in the steppe and the Crimea. Pan- tophagous. The beetles eat sprouting seeds and generative organs of grain (wheat, panicgrass, barley) and medicinal (St John’s wort) crops, and also some wild-growing herbs (Poaceae, Compositae). The larvae are mostly carnivorous, eat eggs of other insects (Sakine and Iskender, 2009). Amara (Bradytus) fulva Muller, 1776. Body size – 7.5–9.5 mm. The pre-scutellum stria of the elytra have no setiferous pore. Is of uniform yellow or yellow-brown color, the upper side has a greenish gloss.The body is wide, the pronotum is about 1.9 times wider than its length. Distributed in Europe, the East Mediterranean, the Eu- ropean part of Russia and Siberia (Niemela and Halme, 1992). Dis- tributed throughout Ukraine. Pantophagous. The beetles sometimes cause damage to sprouting seeds of grain (wheat, barley, maize), technical (flax, mustard) and garden (onion, lettuce) crops, the young shoots of potatoes, seedlings of maple and generative organs of wild-growing herbs (Poaceae, Cyperaceae and others). Amara (Celia) bifrons Gyllenhal, 1810. Body size – 5.5–7.0 mm. The entire pronotum, except the main impressions, is uniformly swollen, its lateral edges are not explanate]. The basis of the pronotum is of almost the same width as the basis of the elytra. Narrow, the posterior angles of the pronotum are blunted, and often the entire base is deeply punctured. Reddish-brown, often with metallic gloss, the antennae are of the same reddish color. Distributed in Europe, North Africa, the European part of Russia, Siberia, Central Asia and in the Near East (Noreika and Kotze, 2012). Distributed throughout Ukraine, damage recorded in the forest-steppe and northern steppe zones. Causes damage to grain (wheat, rye, maize), technical (mustard, thymes), garden (onion, sorrel), medicinal (yarrow (Achillea), 52 Chapter 2. Coleoptera pests of stored food supplies and field crops plantain, St John’s wort, daisies) crops, fallen fruit in gardens, feeding observed on more than 20 species of wild-growing herbs (Poaceae, Polygonaceae, Chenopodioideae, Cruciferae, Euphorbiaceae, Rubia- ceae, Compositae). These beetles are attracted to formalin, which is used as a fixating liquid in traps (Holopainen and Varis, 1986). Amara (Curtonotus) aulica Panzer, 1796. Body size – 11–14 mm. The projection of the prothorax behind the front coxae has no fringing. The pronotum in front of the posterior angles is clearly sinuate. The posterior angles of the pronotum are sharp, protrude to the sides, the lateral edges are rounded almost to the posterior angles. Wide, black-brown, the upper part sometimes has a bronze gloss, the antennae and legs are reddish. Distributed in Europe, the European part of Russia, West Siberia, Central Asia and the Near East, introduced in North America (Eyre, 1994). Distributed throughout Ukraine. Pantophagous. Sporadically harms sprouting seeds, more rarely flowers of grain crops (wheat, corn, rice), grain legumes (peas), technical (cotton) and forage (clover, lucerne) crops, seedlings of maple, hornbeam and some other species of wild-growing herbs (Composi- tae). The beetles are tolerant to high temperatures and are xerophiles (Lambeets et al., 2008). Amara (Curtonotus) convexiuscula Marsham, 1802. Body size – 10–13 mm. The episterna of the metathorax bear dense and usually coarse punctures. The front edge of the pronotum is slightly wider than its base. The body is flattened, the upper part usually has a metal gloss. Distributed in Europe, the West and partly the East Mediterranean, the European part of Russia, Siberia, and the Near East (Eyre, 2004). Distributed throughout Ukraine, except mountain regions of the Carpathians and the Crimea. Pantophagous. Is recorded as a pest of grain (wheat, rye, rice), technical (beet, mustard, fennel (Foeniculum), medicinal (plantain, restharrow) and forage (clover) crops, sprouting seeds of hornbeam and flowers of 7 species of wild-growing herbs (Poaceae, Chenopodioideae). Amara (Xenocelia) ingenua Duftschmid, 1812. Body size – 8.5–10.5 mm. The base of the pronotum is narrower than the base of the elytra, the body is wide and large. The main impressions of the V. I. Rusynov, V. O. Martynov, T. M. Kolombar 53 pronotum are in the form of coarse punctures. The head is flattened, the eyes are slightly protuberant. Jet-brown, the upper part is usually with bronze gloss, the elytra of females are mat; the epipleura, antennae and legs are rusty red. Distributed in Europe, the West and East Mediterranean, the European part of Russia, Siberia, Central Asia and the Near East. Distributed throughout Ukraine (Putchkov, 2011). Pantophagous. Sporadically damages sprouting seeds of grain (wheat, maize), technical (kenaf (Hibiscus cannabinus), cannabis), garden (carrot, lettuce) crops, flesh of strawberry fruit, in fruit gardens - fallen fruit and generative organs of some wild-growing grasses. Amara (Xenocelia) municipalis Duftschmid, 1812. Body size – 5.5–7.0 mm. The elytra have obvious pre-scutellun stria. The antennae are dark, starting from the third segment. The eyes are moderately protuberant. Black, the upper part has bronze or greenish gloss, the epipleura of the elytra are usually brown; the legs are rusty red. Distributed in Europe, the European part of Rus- sia, Siberia and the Near East (Schwerk and Szyszko, 2009). Dis- tributed throughout Ukraine. Pantophagous. Sporadically harms the generative organs of grain (wheat, rye) and technical (mint) crops; feeding observed on 7 species of wild-growing herbs (Poa- ceae, Labiatae, Compositae). Amara (Zezea) chaudoiri Schaum, 1858. Body size – 8–10 mm. The front angles of the pronotum almost or completely do not go beyond the front edge. The legs are completely rusty-red. The upper part is blue or black-blue. Distributed in Europe, the Mediterranean, the European part of Russia, Siberia, Central Asia and the Near East (Saska and Honek, 2003). Distributed throughout Ukraine, except the mountain belt of the Carpathians and Crimea. Pantophage. The beetles eat the generative organs of 28 species of meadow and swampy herbs (Sagittarieae, Poaceae, Cyperaceae, Juncaceae, Cru- ciferae, Chenopodioideae, Compositae) and sporadically grain (wheat, rye), garden (sorrel) and medicinal (poppy, plantain) crops. Amara (Zezea) plebeja Gyllenhal, 1810. Body size – 6.0–7.5 mm. The upper spur of the protibiae is wide, three-denticled. The anten- 54 Chapter 2. Coleoptera pests of stored food supplies and field crops nae are always two-colored: the main 3-4 segments are yellow-reddish, the rest are black. The upper part is metallic. The front edge of the pronotum is sinuate, the front angles are prominent. The groove of the front edge of the pronotum is blue, the front edges are protruding; the posterior angles are flattened, the main pits are densely punctured. The striae of the elytra are thin, not deep. Bronze, with green or brass gloss, the femora are black, the tibiae are reddish. Distributed in Europe, the West and partly the East Mediterranean (except Asia Minor), the European part of Russia, Siberia (Van Huizen, 1977). Distributed throughout Ukraine, except the Crimea. Pantophagous. Causes damage to the generative organs of 21 species of wild-growing herbs (Poaceae, Cruciferae, Euphorbiaceae, Compositae), sometimes grain (maize, wheat, rye, barley), technical (beet, mustard, thymes, flax), garden (cabbage, sorrel), medicinal (poppy, madder (Rubinia tinctorum), plantain, sage, wallflower, daisy, dandelion) and forage (vetches, lupin) crops. Amara consularis Duftschmid, 1812. Body size – 7–9 mm. The body color is jet-black or deep brown, the antennae and the legs are rusty red. The pronotum is no less than 1.7 times wider than its length. The groove in the front of the posterior angles of the pronotum is barely noticeable. The body is wide, the elytra are about 1.3 times longer than their width. The upper part is dark brown, sometimes with light metallic gloss. Distributed in Europe, the European part of Russia, Siberia and Central Asia (Kromp, 1990). Distributed throughout Ukraine. Pantophagous, the beetles sometimes damage sprouting seeds of grain (wheat, rye, barley, maize), technical (beet, mustard), forage (sorghum, clover) crops, maple, hornbeam, berries of strawberry, fallen fruit in gardens. Amara tricuspidata Dejean, 1831. Body size – 7.0–8.5 mm. The front angles of the pronotum are moderately prominent, its basis is not flattened, only the main impressions bear scattered punctures. The striae of the elytra are moderately deep in the front, deepened towards the top. The upper part is dark-bronze without greenish tone, the legs are entirely black or the tibiae are slightly lighter than the femora. Patchily distributed European species (Desender and V. I. Rusynov, V. O. Martynov, T. M. Kolombar 55

Bosmans, 1998). Distributed throughout Ukraine. Pantophagous. In meadows and swamps, eats the generative organs of 18 species of plants (Sagittarieae, Poaceae, Cyperaceae, Polygonaceae, Caryophyl- laceae), in fields - wheat.

Subfamily Pterostichinae

Pterostichus cupreus Linnaeus, 1758. Body size – 10.5–13.5 mm. The third interval of the elytra has one pore. The fringing of the lateral edges of the pronotum gradually enlarges towards the back and is explanate. The first and the second segments of the antennae are red. The shoulder denticle of the elytra is not obvious. The head is densely punctured. The elytra are slightly wider than the base of the pronotum. The upper part is copper-red, bronze, green or black with greenish gloss, rarely blue; sometimes the femora, and rarely the entire legs are red. Distributed in Europe, the European part of Russia, Siberia, Central Asia (Thomas et al., 2001). Distributed throughout Ukraine. Predator. The beetles sometimes damage the sprouting seeds of grain (maize), grain legumes (peas, wild beans, broad beans), technical (beet), garden (lettuce, spinach, radish (Raphanus sativus var. radicula) and forage (clover, lucerne, vetches, lupin) crops, maple, hornbeam, ash, young shoots of potato, flesh of the fruits of strawberry, tomato, fallen fruit in gardens (Brygadyrenko, 2016). Pterostichus melanarius Illiger, 1798. Body size – 12.0–17.5 mm. The claw segment of the feet is covered with setae in the lower part. Uniformly black, glossy, the striae of the elytra are deep. Distributed in Europe, the West and East Mediterranean, Sibe- ria, Central Asia (Fournier and Loreau, 2001). Distributed throughout Ukraine. Predator. The beetles eat seedlings of grain (corn), grain legumes (peas, broad beans, soya beans), technical (beet) and garden (lettuce) crops, sprouting seeds of maple, hornbeam, ash, fruits of blackberry, stone bramble (Rubus saxatilis), tomato and fallen fruit in gardens (Chapman et al., 1999). Zabrus (Pelor) spinipes Fabricius, 1798. Body size – 18–23 mm. The body is thickset, massive, significantly swollen. The pronotum 56 Chapter 2. Coleoptera pests of stored food supplies and field crops is entirely covered in punctures. The elytra have fine puncture rows. Black, slightly glossy, the elytra of females are mat. Distributed in the eastern parts of West Europe, the East Mediterranean, the European part of Russia, Siberia and Central Asia (Gokturk and Celik, 2017). Can be found throughout Ukraine, where three zones of damage are distinguished: heightened - Odessa, Mykolaiv, Kherson Oblasts and steppe zone of Crimea; sporadic damage - Zaporizhia, Kher- son Oblasts and steppe Crimea; insignificant damage - north steppe and south forest-steppe. Damages grain (wheat, rye, maize, barley, panic grass, oats, rice), less commonly technical (beet, sunflower), forage (Sudan grass, timothy, false oat-grass) crops and four species of wild-growing herbs (Sumarokov, 2004). Zabrus (Zabrus) tenebrioides Goeze, 1777. Body size – 14– –16 mm. The pronotum bears dense punctures only on the base. The elytra have deep puncture rows. Jet-black, often with bronze gloss, the antennae and legs are brown-red. Distributed in the middle belt of Europe, the West and East Mediterranean, central and south zones of the European part of Russia, south part of West Siberia and Central Asia (Osborne, 1971). Distributed throughout Ukraine, where three damage zones are distinguished: heightened damage - Odessa, Mykolaiv, Kherson, and the lowland part of the Crimea; moderate damage - central and south regions of Kirovohrad, Dni- propetrovsk and Zaporizhia Oblasts, and also Chernivtsi and Za- karpattia Oblasts; insignificant damage - forest-steppe zone to the south border of Polesie. Damages grain (wheat, rye, maize, panic grass, oats, rice, sorghum, buckwheat), technical (sunflower, fennel), garden (lettuce), forage (Sudan grass, timothy, false oat-grass, fescue (Festuca), orchard grass (Dactylis glomerata), wheatgrass (Agropy- ron) crops and about 10 species of wild-growing grasses. The greatest damage is caused to winter wheat (Bassltt, 1978).

Subfamily Scaritinae

Clivina (Clivina) fossor Linnaeus, 1758. Body size – 5.0–6.5 mm. The body is elongated, jet-black or reddish with no metallic gloss. V. I. Rusynov, V. O. Martynov, T. M. Kolombar 57

The pronotum has no impressions on the sides from the middle line. The terminal sternum of the abdomen has fine microsculpture. The pronotum and the elytra are of one color, jet-black or brown (reddish among incompletely colored individuals). Distributed in Europe, Mediterranean, Central Asia, Siberia, introduced in North America and distributed across most of the USA and Canada (Nel- son and Reynolds, 1987). Distributed throughout Ukraine. Preda- tor. Adult individuals are able to fly, which allows them to spread (Gerken et al., 1991). The beetles, in some cases and the larvae, gnaw out the seedlings and underground parts of grain (maize), technical (beet, mustard, cotton plant), garden (cucumber, cabbage, lettuce, onion) and medicinal (Valeriana) crops.

Subfamily Trechinae

Bembidion assimile Gyllenhal, 1810. Body size – 2.8–3.5 mm. Piceous black, the front part of the body is greenish, the elytra have a bluish tone. Pronotum dull from dense, reticulate microsculpture assimile with elytral striae deeper and more coarsely punctate. An- tennae and legs stouter. Elytra with pale apex and preapical macula but in anterior half without or with indistinct spots. Wings often reduced (Lindroth, 1974). Distributed in Europe, the Mediterranean, North America, Siberia and Central Asia (Zulka, 1994). Distributed throughout Ukraine. Predator. The beetles sometimes eat the generative organs of meadow grasses. Bembidion lampros Herbst, 1784. Body size – 3.0–4.5 mm. Up- per surface with metallic, usually brassy, rarely bluish, lustre. Base of antennae (at least first segment underneath) and legs reddish, but femora and tarsi often infuscated. Frontal furrows somewhat dilated at middle. Seventh elytral stria lacking or consisting of a row of weak punctures anteriorly. Penis with external left sidefold (Lindroth, 1974). Distributed in Europe, the West and East Mediterranean, North America, Siberia and Central Asia (Wallin and and Ekbom, 1988). Distributed throughout Ukraine. Predator. Sometimes damages the seedlings of grain (wheat), technical (flax) and garden (tur- 58 Chapter 2. Coleoptera pests of stored food supplies and field crops nip rape) crops, cabbage shoots, flesh of strawberry fruits, sprouting seeds of pine, spruce, maple. Bembidion quadrimaculatum Linnaeus, 1758. Body size – 2.8–5 mm. A small species with very long legs. Black, forebody more or less metallic, elytra with humeral and almost constantly with preapical spot yellow, and often apex, sometimes also suture brown. Four basal antennal segments and legs rufo-testaceous (or femora slightly infuscated) (Lindroth, 1974). Distributed in Europe, North America, the European part of Russia, Siberia, Central Asia. Distributed throughout Ukraine. Predator (Grafius and Warner, 1989). The beetles rarely harm the generative organs of grain (wheat, rye), medicinal (plantain, poppy) and seedlings of garden (onion, radish (Raphanus sativus var. radicula) crops, cabbage plants, flesh of strawberry fruits, fallen fruit in gardens and sprouting seeds of spruce and pine.

Suborder Polyphaga

Family Anobiidae

Subfamily Anobiinae Stegobium paniceum Linnaeus, 1758. Body size – 2–3 mm. The capitulum of the antennae of males is prominent, of females slightly longer than other antenna segments together, notably larger than the segments of the flagella. Red-brown or rusty-yellow; the body has dual hair cover – protruding and appressed. The elytra are less than two times longer than their width. Distributed globally. Significant pest of various food products and materials of mostly plant origin (Lucas and Riudavets, 2002). The adult beetles do not feed. Female lays 20-60 eggs on different products and goods, which the larvae feed on. The larvae penetrate the layer of food substrate and dig tunnels in it, which by the end of the larval development are about 2 mm in diameter (Kučerová and Stejskal, 2010). Before pupation, the larva creates a pupa cradle and fills it inside with wood flour. The beetle gnaws out a flying hole of 0.9 to 1.6 mm in diameter, and flies out. The beetles often gather on windows and window panes. V. I. Rusynov, V. O. Martynov, T. M. Kolombar 59

Larvae cause damage to grain, bread, various flour products, many domestic objects, wicker baskets, armchairs, pharmaceutical products, plywood products and many other objects. The duration of the larva’s development depends on the environmental temperature. At the temperature of 17 °С, development lasts 37 days, 24 – 17 °С, and at 28 °С, it is completed in 8 days. Therefore, in different climatic conditions, the number of generations is different. During one year in Europe, usually one generation develops, and in the southern states of the USA - up to four. At a temperature of 4-5 °С, development stops, and the larvae enter diapause which sometimes lasts up to four months and often ends in their death (Jian et al., 2010).

Subfamily Ptininae Niptus holosericeus Faldermann, 1836. Body size – 4.0–4.5 mm. The pronotum is almost ball-shaped, the middle line is distinguished weakly and only in the posterior part. The elytra are ball-shaped, puncture furrows (striae) are very insignificant. The entire body is covered with hairs of golden color, which hide the main surface. Distributed in Europe, the Mediterranean, North Africa, the European part of Rus- sia and the Near East (Manachini, 2015). Can be found throughout Ukraine. Synanthrope. Usually feeds on different organic remains, can damage rugs, leather goods and grain products. Epauloecus unicolor Piller & Mitterpacher, 1783. Length – 2.2–2.8 mm.; width – 1.3–1.5 mm. Body form much as in N. holo- leucus Fald. but with the pronotal disk less convex and with the sub- basal angles before the prothoracic constriction less broadly rounded. Integuments reddish-brown. Vestiture consisting of rather coarse, golden hairs. Head very finely and densely alutaceous, opaque, the sculpture almost concealed by the closely appressed hairs. Antennae two-thirds as long as the body, the intermediate segments only slightly longer than wide. Pronotum shining, roughly and very densely punctate, the sculpture somewhat obscured by the hairs; the latter close, decumbent; some of them, on each side of the median line and on each side before the constriction, semi-erect, this usually causing the median line to appear sulcate and the sub-basal angles to appear tu- 60 Chapter 2. Coleoptera pests of stored food supplies and field crops mid. Elytra shining; the sides strongly rounded; the striae represented by rows of deep, coarse punctures behind each of which is a prostrate hair; intervals subequal in width to the striae, each supplied with two or three rows of prostrate hairs and a single row of semi-erect hairs; all hairs of the elytra equal in length, the length of each subequal to the width of an interval (Norman, 1875). Distributed in Europe, Mediterranean, Near East, the European part of Russia. Can be found throughout Ukraine. Usually feeds on organic remains of plant origin. Ptinus (Ptinus) fur Linnaeus, 1758. Body size – 2.0–4.3 mm. The penultimate segment of the legs is simple (not bifurcated) in both sexes. The hairs on the elytra of females in the striae and in the intervals between them are long, protrude; in males, the hairs in the striae and intervals are appressed and of the same length. The elytra have white scales. The pronotum of both sexes bears two longitudinal, usually narrow, spots of yellow hairs on the disk, which stretch to its middle. The size and color vary, but the spots described are always present. Prefers humid habitat, can be found in decomposing wood and other plant detritus, sometimes on the remains of animals. Distributed throughout the globe (Howe and Burges, 1951). Can be found throughout Ukraine. Females are flightless, the wings are developed only in males. Due to specific body structure, Ptinidae beetles are often mistaken for spiders. Beetles and larvae damage various products and materials: rusks, groats, flour, grain, hay, taxidermy mounts, insect collections of. Resistant to heightened and low temperatures. Ptinus (Ptinus) latro Fabricius, 1775. Body size – 3–4 mm. The elytra of males and females are different in shape, but the females have no obvious shoulders, the elytra of males are narrow and parallel. Raised hairs on the body are yellow. The elytra of females are long and elliptical, with thin puncture furrows, their intervals bear a row of similar protruding setae, among males - stretched and narrow, the intervals significantly wider than the striae. The punctures on the apical part of the second stria of the elytra are usually sepa- rated from one another by intervals which are 1.5–2.0 times larger than the puncture in diameter. The punctures on the metathorax are 2–3 times as long as their width. Color is dark. Distributed throughout V. I. Rusynov, V. O. Martynov, T. M. Kolombar 61 the Palearctic, except the north. Can be found throughout Ukraine. Damages stored grain, flour, raw tobacco (Stejskal et al., 2011). Ptinus (Ptinus) pusillus Sturm, 1837. Body size – 1.8–3.0 mm. The pronotum bears no longitudinal spots of appressed yellow hairs. Meso- and metatibiae of males have one thin apical spur. The antennae of females are thin, shorter than half of the body, their 4-10 segments no more than two times longer than their width on the top. The pronotum has only poorly developed hair patches. The front sash on the elytra is straight, enlarged on the sides. The elytra of males are stretched, but their sides are not parallel, moderately rounded. Dis- tributed throughout the Palearctic, uncommon (Smith and Barker, 1987). Damages stored flour and grain. Ptinus (Cyphoderes) raptor Sturm, 1837. Body size – 3–4 mm. The pronotum on both sides of the disk has highly lifted light small “pillows” of hair. Small “pillows” of hair on the pronotum are small, stretch to the front no farther than the middle of the pronotum, the same in both sexes, divided by a wide furrow. The pronotum has no obvious ansiform hair patches on the sides. The elytra of females bear moderately long, protruding hairs, appressed in males, two white sashes nearly reach the elytral suture. Distributed throughout the globe. In Ukraine, can be found within the forest and forest-steppe zones (Brown, 1940). Is different from P. f ur by presence of two yellow felt “pillows” on the pronotum. Causes damage to grain stores. There are cases of damage to honeycombs. Ptinus (Bruchoptinus) rufipes Olivier, 1790. Body size – 3.5 mm.The base of the pronotum has no raised longitudinal tubercle on each side. The pronotum has no sharply demarcated small highly raised “pillows” of hair, but sometimes has ridges of hair. Its disk does or does not bear small brush like patches of hairs. The sexual dimorphism is manifested sharply: male is narrow with long, almost parallel, elytra, whereas the elytra of females are oval or elliptical and wide. The penultimate segment of the legs of males and females is not sharply, but obviously bilobate. The elytra of males are significantly stretched, covered in yellowish stripes of same color, with no light sashes. The elytra of females are significantly swollen, 62 Chapter 2. Coleoptera pests of stored food supplies and field crops oviform, with two transversal zigzag sashes, of which the posterior one ends on the elytral suture, and bears small spots at the apices. The main color is black or black-brown (Howe, 1959). Distributed in Europe, in the territory of the former USSR. Can be found throughout Ukraine, mostly in the forest and forest-steppe zones. Has been found under bark of trees in natural conditions and in grain stores. Damages grain and grain products.

Family Bostrichidae

Subfamily Dinoderinae Rhyzopertha dominica Fabricius, 1792. Body size – 2.5–3.0 mm. Oblong, reddish-brown, glossy. The front part of the pronotum has small, flatly pointed teeth, positioned as regular semicircles. The rows of punctures are coarse. The larvae are polyphagous, and harm not only products of plant origin, but also of animal origin, for example leather (Breese, 1960; Edde, 2012). Distributed in the Afro- tropical region, Australia, the Azores, the Balearic Islands, Belarus, Great Britain, the Canary Islands, Corsica, Croatia, Cyprus, Den- mark, the Dodecanese Islands, France, Germany, Greece, Ireland, Italy, Latvia, the Near East, the Nearctic region, the Neotropical region, North Africa, the Oriental region, Poland, Portugal, Romania, Sardinia, Sicily, Spain, Austria, Belgium, the Czech Republic, Esto- nia, Finland, Slovakia, Sweden, Netherlands. Globally distributed (Fields et al., 1993). Colonised port cities of the USSR, including Ukraine. Damages grain and different groats.

Family Dermestidae

Anthrenus (Florilinus) museorum Linnaeus, 1761. Body size – 2.2–3.6 mm. The body is densely covered in scales. The antennae comprise 8 segments, with the capitulum of 2-segments. The sockets for folding in the antennae are very deep. The epimera are not seen. Larvae live in nests of birds, wasps and bees. The beetles are often found on flowers of meadow and steppe plants. The V. I. Rusynov, V. O. Martynov, T. M. Kolombar 63 upper part is covered in black scales with yellow-brownish scales; white and yellow-brown scales cover the posterior angles and the base of the pronotum and form three thin tortuous sashes on the elytra. Distributed in Europe, North America, the European part of Russia, Siberia and Central Asia (Woodroffe and Southgate, 1954). Can be found throughout Ukraine. Damages zoological, entomological collections, food supplies and food material. The beetles feed on nectar and pollen of umbellifers, common in human habitations. Anthrenus (Anthrenus) scrophulariae Linnaeus, 1758. Body size – 3.0–4.5 mm. The elytra bear black scales with three narrow sashes of white scales, interrupted at the elytral suture. The elytral suture has red and yellow scales. Sometimes the entire elytra bear spots of black, white and yellow scales. Dis- tributed in Europe, Asia, the USA, the European part of Rus- sia (Hasan et al., 2008). Can be found throughout Ukraine. The beetles appear in spring and in early summer on flowers and fruit trees and on different crucifers. In mid-June, they damage mustard, courgettes, wild mustard (Sinapis arvensis), damaging flowers and leaves. Harm leaves of apricot and peach. Can be found in bird nests and in houses, damages wool products, zoological and entomological collections. Anthrenus (Florilinus) verbasci Linnaeus, 1767. Body size – 1.7–3.2 mm. Internal edge of the eye has no groove. The upper part is in covered with elongated jet-black and yellow scales. The sides of the pronotum, its base and three sinuous thin sashes on the elytra bear white scales. Distributed throughout the Palearctic (Woodroffe and Southgate, 1954). Can be found throughout Ukraine, in bird nests and in houses. Damages woolen products, zoological and entomological collections. Dermestes (Dermestes) lardarius Linnaeus, 1758. Body size – 7.0–9.5 mm. The body is comparatively large. The forehead has no median ocellus. The antennae comprise 11 segments, with a 3-segment capitulum. Males usually have a little pit with a brush of long hairs. The pronotum is covered with black hairs with spots 64 Chapter 2. Coleoptera pests of stored food supplies and field crops of yellow hairs. The sash on the elytra bears yellow-grey hairs with 6 round spots of black hairs (Kingsolver, 1991). Distributed in Eu- rope, North America, the European part of Russia and the Near East. Can be found throughout Ukraine, in bird nests and in houses. Damages leather and fur goods, meat and fish products. The larvae attack nestlings of poultry. Trogoderma granarium Everts, 1898. Body size – 1.8–3.0 mm. The antennae consist of 11 segments (rarely 9-10 segments), with 3– –9 segment head. The epipleura of the pronotum have sharply des- ignated pits for the capitulum of the antennae. The tibiae have no spinescent structures. The cuticle is light-red-brown on the top, sometimes the head, pronotum and several spots on the elytra have a darker color. The upper part is covered with yellow hairs, dark parts of the cuticle are covered with light-brown hairs. The antennae usually comprise 11 segments, sometimes 9-10 segments. The antennae head of males consists of 3-5 segments. Distributed in the Afrotropi- cal region, Oceania, North Africa and in the Near East. With grain, it was introduced to Europe, Australia, North America (Banks, 1977). For Ukraine - it is an object of external quarantine. Significant pest in grain stores. The larvae damage long term stores of grain, especially barley, corn, rice, wheat, seeds of cotton plants and other products rich in protein. Can feed on products of animal origin (Jood et al., 1996). Trogoderma versicolor Creutzer, 1799. Body size – 2–5 mm. The cuticle of the elytra is black or brown-black, with three reddish- brown sashes and several spots. The upper part is covered with black or dark-brown hairs, light areas of the cuticle are covered with white and yellow hairs. Distributed in Europe, North Africa, Western Asia, the European part of Russia, Central Asia, Trans- caucasia, Siberia (Strong et al., 1959). Can be found throughout Ukraine. Damages books, leather, zoological and botanical collections, sometimes grain of maize in grain storages, products of animal origin. Causes damage to beekeeping and sericulture, larvae harms cocoons, silkworms and pupae of the domestic silkmoth (Hadaway, 1956). V. I. Rusynov, V. O. Martynov, T. M. Kolombar 65 Family Cerambycidae

Subfamily Lamiinae Dorcadion holosericeum Krynicky, 1832. Body size – 11.5–17.0 mm. The elytra bear two dark velvety stripes in intercos- tal sections; has marginal, shoulder and the elytral suture stripes, and no light stripe on the spine. The dorsal surface is grey-black or brown, the antennae and legs are black. Distributed in the forest- steppe and steppe zones of Poland, Russia, Belarus, Moldova, Ro- mania, Slovakia, Georgia and Ukraine (Dascălu, 2012). The larvae live in soil, feed on the roots of herbs, mainly grasses (Özdikmen, 2010). Dorcadion carinatum Pallas, 1771. Body size – 15–23 mm. Front legs with large, significantly enlarged segments. The main segments of the antennae are mat, with noticeable fine covering. The front part of the marginal stripe of the elytra usually bears distinct punctures. Has no distinctive white elytral suture. The elytra are significantly elongated.The pronotum bears large non-uniform puncturing. The humerus of the elytra is blunted only in the third apex part. Black, glossy (Özdikmen, 2010). Distributed in forest-steppe and steppe of Russia, Ciscaucasia, Azerbaijan, Kazakhstan, Georgia and Ukraine. Feeds on leaves of grasses (Biliavskyi et al., 2012).

Family Chrysomelidae

Subfamily Chrysomelinae Chrysolina (Chrysomorpha) cerealis Linnaeus, 1767. Body size – 6–11 mm. The pronotum has rounded sides; the puncturing on the elytra is chaotic. The pronotum has a notable fold along the side edges in the base part. Green, with blue, bronze, violet or purple longitudal stripes. Distributed in Central and Southern Eu- rope, the European part of Russia, the Caucasus, Central Asia and Siberia (Bieńkowski, 2013). Can be found throughout Ukraine. Beetles and larvae are seen on the leaves of cereals from May to September (Marshall, 1979). 66 Chapter 2. Coleoptera pests of stored food supplies and field crops

Gastrophysa (Gastrophysa) polygoni Linnaeus, 1758. Body size – 4–5 mm. The pronotum, legs and the bases of the antennae are orange- red, the head and the elytra are blue, violet or blue-green. Distributed in West Europe, North America, North China, Mongolia, Korea, the European part of Russia, Transcaucasia, Central Asia and Siberia. Can be found throughout Ukraine (Deroe and Pasteels, 1982; Bieńkowski, 2013).The beetles and larvae feed on different species of wild-growing Polygonaceae, but often damage the leaves of buckwheat, lucerne, vetches, beet, mustard, grain crops and other field crops.

Subfamily Criocerinae Oulema melanopus (Linnaeus, 1758). Body – 4.6–5.3 mm. Head, elytra, coxae, trochanters, tarsi, and tibial apices black; pronotum, femora, and most of tibiae orange to red orange; all black parts of body (except outer 8 antennal segments, apices of tibia, and tarsi) also with a distinct bluish reflection; surfaces shiny. Anterior and basal margins of pronotum may be narrowly black; sometimes elytral reflection greenish; mandibles often reddish. Head .The antenna clearly more than 1/2 length of body; outer segments elongated, segments 7 and 11 each about 2 times as long as wide, segments 8–10 each less than 2 times as long as wide. Vertex with a distinct, median groove; surface weakly convex to flat and densely punctate, punctures variable in size. Clypeal area with dense punctation and with setae obscuring surface. Pronotum. Width and length subequal, widest medially; constriction located at basal 1/4, width at constriction 88% of medial width; medially at basal 1/4, with or without a small fovea; large, not dense, irregular punctures at sides extending anteriorly to middle, on disk extending to near base; at basal 1/4, a coarsely punctate transverse depression; surface also with sparse, very fine punctures. Elytra. At basal 1/4, a weak to no depression; intervals between striae with weak development of fine punctures and transverse grooves; infrequently in basal 1/4, large punctures of striae transversely aligned; 3d stria with 13-18 punctures in basal 1/2 of elytron; 9th stria complete. Elytra much elongated, each elytron 3.5–3.7 times as long as wide. Ventral Surface. Metastemum with large punctures, moderate in density, a V. I. Rusynov, V. O. Martynov, T. M. Kolombar 67 seta arising from each puncture; large punctures nearly absent medially except along anterior margin; surface also with fine punctures. Abdomen finely scabrous, especially at sides; stemites 1–4 below apex with transverse series of moderate-sized punctures; stemite 5 with scattered moderate-sized punctures. Length. Aedeagus. In lateral view widest medially, widest point behind declivity, apical tip pointed downward and sharply pointed, lower apical margin broadly sinuate, upper apical margin nearly flat and curved downward apically; in dorsal view sides subparallel basally but narrowed anteriorly, small orifice overlaid by 3 lobes, 2 lateral lobes broadened and concealing median lobe, portion behind orifice flat; internal processes simple, in dorsal view symmetrical but of no distinctive form (White, 1993). Distrib- uted in Europe, the Mediterranean, North America, North Africa, the Near East, Asia Minor, Mongolia, the European part of Russia, the Caucasus, Siberia, Central Asia. Can be found throughout Ukraine, more abundant in the steppe, central and eastern forest-steppe. Dam- ages oats, barley, durum wheat, corn, panic grass (Kher et al., 2012).

Subfamily Eumolpinae Pachnephorus (Pachnephorus) tessellatus Duftschmid, 1825. Body size – 2.5–3.5 mm. The upper part is densely covered with grey scales which hide the main body surface. Distributed in Mongolia, Albania, Austria, Bosnia and Herzegovina, Bulgaria, the Czech Re- public, Estonia, France, Germany, Greece, Hungary, Italy, Latvia, Lithuania, the Near East, Poland, Romania, Slovakia, Spain, Swit- zerland, the European part of Russia, the Caucasus, Transcaucasia, Central Asia and Siberia (Gavrilović and Ćurčić, 2011; Morse et al., 2014). Can be found in the forest-steppe zone of Ukraine. The beetles sometimes damage the seedlings of winter wheat, and in West- ern Siberia were recorded to damage sunflowers.

Subfamily Galerucinae Chaetocnema aridula Gyllenhal, 1827. Body size – 2.0–2.5 mm. The upper part is bronze or copper-bronze. The puncturing on the elytra is randomly patterned, organized in rows in some places. Prolonged- 68 Chapter 2. Coleoptera pests of stored food supplies and field crops oval, dark-bronze; the punctures on the elytra are quite small on the top and dense. The shoulder tubers are swollen. Distributed in West Europe, the European part of Russia, the Caucasus, Transcaucasia, Central Asia and Siberia (Schmitt and Rönn, 2011). Can be found throughout Ukraine. Harms winter and summer wheat, barley, rye, oat, Se- taria italic subsp. italica, wheatgrass, fescue and other grasses (Kaplin and Antonov, 2006). Galeruca (Galeruca) pomonae Scopoli, 1763. Body size – 6– 12 mm. The side margin of the elytra ends on the shoulders or near the posterior angles of the pronotum. The lateral stria of the pronotum is significantly impressed and remote from the side edge; each elytron bears four initial ribs, with one or few incomplete secondary ribs. The upper part is yellow-brown, rarely black. Distributed in Europe, North America (introduced), Northern Iran, the European part of Russia, Siberia, the Caucasus and Kazakhstan (Schmitt and Rönn, 2011). Can be found throughout Ukraine. Beetles and larvae sometimes damage cabbage, carrots, sugar beet, rapeseed, winter and summer cereals, and also field and garden crops. Phyllotreta vittula Redtenbacher, 1849. Body size – 1.8–2.3 mm. The forehead and much of the vertex is covered in punctures. The internal edge of the yellow stripe of the elytra is straight and near the apex is slightly creased towards the elytral suture; the head and the pronotum are metal-green. Distributed in Europe, North Africa, Central Asia, Ko- rea, the European part of Russia, the Caucasus, Transcaucasia, Central Asia and Siberia (Schmitt and Rönn, 2011). Can be found throughout Ukraine. Damages wheat, barley, rye, panic grass, maize, Setaria italic subsp. italica, false oat grass, foxtail (Setaria), fescue, meadow-grass (Poa), orchard grass, and other grasses, rarely sugar beet, Cruciferae, an- ise (Pimpinella anisum), mahaleb cherry (Prunus mahaleb) (Vig, 1998).

Family Cleridae

Subfamily Korynetinae Necrobia violacea Linnaeus, 1758. Body size – 4.0–4.5 mm. The upper part is completely blue or green. The antennae and legs are V. I. Rusynov, V. O. Martynov, T. M. Kolombar 69 dark. The elytra have deep puncture rows which gradually disap- pear near the apex. Cosmopolite. Distributed in Afrotropical, Ne- arctic, Neotropical and Oriental regions, Oceania, the Near East, North Africa. The beetles and larvae are predators. Can harm products of animal origin and grain in storages (Özdemir and Sert, 2009).

Subfamily Trogositinae Tenebrioides mauritanicus Linnaeus, 1758. Body size – 6–10 mm. The body is flat with large head and transversal-heart- shaped pronotum. Brown-black. Cosmopolite. Distributed in Af- rotropical, Nearctic, Neotropical and Oriental regions, Oceania. Considered native to Africa. Can be found throughout Ukraine (Sumarokov, 2004). Lives in mills, groat and baking factories, stores, in residential buildings, on ships; in natural conditions - under the bark of diseased and dried out trees. The beetles are predators, can be found under the bark of drying trees, and also in grain stores, where they eat larvae and pupae of different insects. The larvae also consume insects in grain stores, granaries, at the same time they also cause significant damage to grain and grain products (wheat, barley, corn, oats, seeds of cotton plants), dried fruits, vegetables, etc). Prefers flour. Damaged flour becomes dark and obtains unpleasant smell and taste. Over an hour, a single larva harms 1-5 grains, mostly their embryo parts, causing loss of germination. Especially damages grain with moisture over 10 %.

Family Laemophloeidae

Cryptolestes ferrugineus Stephens, 1831. Body size – 1.5–2.2 mm. Clearly distinguished from others by impressed lines on the sides of the pronotum. The body is flat, males have wider head and longer antennae. The width of the pronotum is slightly greater than its length. The antennae of males reach the middle of the body, the antennae of females do not. Rusty red. Cosmopolite. Distributed in Austria, Belgium, Great Britain, Croatia, the Czech Republic, Denmark, Eu- 70 Chapter 2. Coleoptera pests of stored food supplies and field crops ropean part of Turkey, Finland, France, Germany, Greece, Hungary, Italy, Poland, Portugal, Sicily, Spain, Sweden, southern regions of Russia (Hagstrum, 1989; Flinn and Hagstrum, 1998). Can be found throughout Ukraine. Damages stored grain products, similarly to sawtoothed grain beetle (Oryzaephilus surinamensis). Differs from it by smaller size (1.5–2.2 mm). Usually lives in stored grain, sometimes forms large colonies in the breeding grounds with high temperature (30–32 °С) (Campbell and Sinha, 1976). Placonotus testaceus Fabricius, 1787. Body size – 1.5–2.5 mm. The pronotum is almost square, the front angles are stretched into a sharp spike. The antennae are of the same length as the body. Brown-yellow. Distributed in Austria, Belgium, Great Britain, Cro- atia, the Czech Republic, Denmark, European part of Turkey, Fin- land, France, Germany, Greece, Hungary, Italy, Poland, Portugal, Sicily, Spain, Sweden, central and southern belts of Russia, in the Far East. Can be found throughout Ukraine (Bondarenko, 2013). Feeds on flour and groats of heightened moisture. Sometimes gnaws out the sieves in flour mills. The development of a generation lasts around three months. Does not develop in substrate at moisture below 15–18 %.

Family Nitidulidae

Subfamily Carpophilidae Carpophilus hemipterus Linnaeus, 1758. Body size – 2–4 mm. The body is wide, oval, swollen on the top; the length of the elytra equals their total width. Black or brown, the elytra have a yellow spot on a shoulder tubercle and a large spot behind the middle. Cos- mopolite. Distributed in the Afrotropical, Nearctic, Neotropical and Oriental regions, Oceania, the Near East, North Africa (Dowd and Bartelt, 1991). Polyphagous storage pest. Beetles and larvae feed on different products, especially dried fruits and baked bread, mainly white (Dobson, 1954). Harms also wheat, barley, flour, peanuts, lettuce seeds , etc. V. I. Rusynov, V. O. Martynov, T. M. Kolombar 71 Family Silvanidae

Oryzaephilus mercator Fauvel, 1889. Body size – 3.0–3.5 mm. The beetles are red-brown or brown. The temples are three times shorter than the eyes. The clypeus of males has no horns, and the femora - no teeth. Distributed in Austria, Belarus, Belgium, Great Britain, Bulgaria, Corsica, Croatia, the Czech Republic, Denmark, European part of Turkey, France, Germany, Greece, Hungary, Italy, Lithuania, Poland, Portugal, Romania, Sardinia, Spain, Switzerland (Loschiavo and Smith, 1970). Can be found throughout Ukraine, along with the sawtoothed grain beetle, but much rarer (Allotey and Kumar, 1988). Oryzaephilus surinamensis Linnaeus, 1758. Body size – 3.0– 3.5 mm. The beetles are red-brown or brown. The temples are of the same length as eyes. The sides of the males’ clypeus are often deflected as horns, and the metafemora in the lower part behind the middle have a small tooth. Cosmopolite. Distributed in Europe, the Mediterranean, the European part of Russia, except northern regions (Trematerra, 2000; Trematerra and Sciarretta, 2004). Can be found throughout Ukraine. Damages embryos of grasses, the disks of sunflowers, flour, dried fruits. Beetles and larvae feed on cracked grain and products of grain processing, rice, dried fruits, dry meat, medical plants, etc. Does not damage wholegrains. The beetles can live without food for a long time. Lives in grain stores, on confectionery and pasta factories, in stores, flour mills, groat and compound feed factories (Fleming, 1988).

Family Curculionidae

Subfamily Cossoninae Caulophilus oryzae (Gyllenhal, 1838). Body size – 2.5–4.0 mm. Beetles vary in size from reddish-brown to jet-black with slight gloss. The tubular extension is not enlarged in the upper part in the form of a spade. The eyes are swollen. The flagellum of the antennae comprises seven segments. The pronotum is uniformly covered in punctures of small rounded spots. The elytra have deep 72 Chapter 2. Coleoptera pests of stored food supplies and field crops puncture rows. Differs from related species of storage weevils by wide and short tubular extension of the head. Caulophilus oryzae or broad-nosed grain weevil has the proportion of the tubular extension to the length of the pronotum and elytra as 1 : 1 : 2, whereas other storage weevils – 1 : 1 : 1. Also, the femora of all legs of the broad-nosed grain weevil are thickened from the middle to the apex; the protibiae have a groove on the internal side (Mordkovich and So- kolov, 1999). The egg is 0.4-0.5 mm long, white, transparent, oblong (Migulin, 1983). The larva is white or yellowish. The body is slightly crescent-bent, wrinkled, bears rare hairs, has no legs. The length is 2.5–3.0 mm. The length of the pupa is 2.8-3.0 mm, the width is around 1.3 mm, white in the beginning and afterwards turns yellow. The tibiae have two short setae, whereas wheat weevil ( Sitophilus granarius) have only one. Winters in stores. The female gnaws a hollow in a seed or other food substrate, where the egg is deposited, and closes the hole with a plug of solidifying secretions. Female lays 200–300 eggs. The egg phase, depending on temperature and moisture, lasts 4 to 14 days. The larva phase lasts around 3 weeks. During development, larva feeds on the internal content of the food substrate. Pupation takes place inside the damaged material (Shamonin, 1986). The pupa phase lasts around 5 days. In optimum conditions, the development cycle is completed in 30 days (Savotikov and Smetnik, 1995). In less favorable conditions, it can last up to 50 days (Eremenko et al., 1967). The beetles are highly resistant to unfavorable conditions: at temperature of 16.6°С, they can exist without food up to 50 days (Migulin, 1983). Distributed in countries of North and Central America; introduced in countries of West- ern Europe and Morocco. Quarantine object for DPRK, Mongolia, Turkmenistan, Kyrgyzstan, Ukraine, Bulgaria, and Romania. Dam- ages seeds of different grain crops, and also chestnuts, acorns, fallen figs, peas, flour, macaroni, etc. Both beetles and larvae are pests . They eat out the internal content of the seed similarly to the granary weevil (Sitophilus granarius) and rice weevil (Sitophilus oryzae). The beetles cannot consume dry solid grain, but can eat moistened, and fractured grain or grain damaged by other pests and lay eggs inside the grains (Sokolov, 2004). Spreads in all phases of the development with fruit, V. I. Rusynov, V. O. Martynov, T. M. Kolombar 73 seeds, grain and other products, which it damages. The beetles fly and can inhabit and damage grain of maize in the field. From there, they become transferred to the storage together with the collected grain (Savotikov and Smetnik, 1995).

Subfamily Dryophthorinae

Sitophilus granarius Linnaeus, 1758. Wheat or granary weevil .Body size – 2.3–4.5, significantly varies (depending on the food which it developed on). The body is narrow, cylindrical, glossy. Young beetles are of light-brown color, adults are almost black. The small head is stretched into a thin tubular extension, the end of which bears the mouthparts of gnawing-chewing type. The antennae bent at a certain angle. The pronotum bears coarse longitudinal pits, the forewings have deep longitudinal striae, and are concrescent. The membraneous hindwings are not developed – the beetle is not able to fly (Sokolov, 2004). The female abdomen is straight in profile; thin anal hole is in the form of transversal slit. The last segments of the abdomen of the male are curved downwards and the anal hole is in the form of curved slit. The laid egg is at first transversal, dirty-white, of regular elliptical shape; over time, this regular shape disappears. The length is 0.6–0.75 mm. Width – 0.3– 0.4 mm. The larva has no legs, fleshy, vermiform, around 3–4 mm long, with brownish head. In shape, the pupa resembles an adult beetle, transparent, 3–5 mm long (Yakovlev, 1974). Female gnaws out a hole in the grain near the embryo, at the bottom of which an egg is placed. The female’s fertility is around 200–250 eggs. For preventing eggs from drying out and protecting them from predators, the egg is covered in slime which solidifies in the air. Few days later, a legless white larva comes out of the egg, shortened, with an outward swelling spine and brown head. Right after hatching, the larva gnaws into the grain, where it spends all of its life, consuming almost all its content. As the larvae complete development, they form a cradle, in which they turn into pupae, which resemble the adult beetle in shape. The pupal development lasts 7–22 days (depending on the air tem- 74 Chapter 2. Coleoptera pests of stored food supplies and field crops perature). After solidifying of the cover, which takes 2–6 days, the beetles of new generation gnaw a rounded hole in the grain and emerge. They feed by gnawing the softest parts of grain, thus damaging a large amount of grain during their lifetime. The beetle avoids illuminated places. After even a slightest disturbance, it falls in stupor, closely appresses the antennae and the legs to the body. Optimum conditions for the weevil’s development in grain is grain moisture 14–16 %, air humidity – 75–95 % and temperature around 25 °С. The duration of the pest’s development from the moment of laying eggs to the imago stage depends on the temperature and moisture. At 17 °С, the development lasts around 80 days, at 20 °С – 70 days, at 25 °С – 34 days and at 28 °С – 1 month. The beetle has a long life. At room temperature and with availability of food, the beetle can live over a year (at temperature of 10–12 °С – 28 months) (Os- molovskiy and Bondarenko, 1980). At the temperature of 5–10 °С, the beetles stop eating, and at 3 °С they fall into chill-coma; at temperature below 0 °С,the weevils gradually die (Sokolov, 2004). The deficiency of moisture slows the development of the weevil, and 11 % moisture is lethal for it. In southern regions, in the conditions of grain stores, the granary weevil can have 2–3 generations, compared to 1–2 in central regions. Beetles, larvae and pupae winter inside grains. The beetles can winter also in cracks in the floor, walls, in cellars and other similar places. Is distributed with all types of infested goods. Especially often with storage equipment, grain-processing machines not cleaned from the remains of old grain, and with undestroyed sweepings and grain wastes unsuitable for usage. Cosmopolite, one of the most harmful and common grain pests. While feeding, the imago harms different grain and processed grain products. Larvae can develop in grain of wheat, rye, barley, oat, rice, maize, buckwheat, panic grass, sometimes in macaroni products and in compressed flour. Grain damaged by the weevil becomes easily accessible for secondary grain pest species – insects and Acari. When the weevil is numerous, the damaged grain becomes unsuitable for human consumption and causes digestive disorders (Sokolov, 2004). Significantly infested V. I. Rusynov, V. O. Martynov, T. M. Kolombar 75 grain becomes hygroscopic and further undergoes self-heating and decomposition. Sitophilus oryzae Linnaeus, 1763. Body size – 2.5–3.5 mm. It looks very similar to S. granarius, differs from it by smaller and thinner tubular extension. The body is brown, mat or slightly glossy; the pronotum bears large dense punctures; each elytron has two red spots; the wings are developed, the beetles are able to fly well (Sokolov, 2004). The elytra are densely covered in puncture striae, and narrow intervals between them are covered in short rows of punctures. The entire pronotum is densely covered in rounded punctures, so in the middle there is no smooth narrow longitudinal line free from these punctures, such as the granary weevil has. All development stages (egg, larva, pupa) in shape and size are similar to the corresponding stages of S. granarius. Larvae are white, fleshy, 2.5–3.0 mm long (Volkov et al.,1955). The pupae are at first white and later become yellowish, up to 2.75 mm (Osmolovskiy and Bondarenko, 1980). The duration of the beetles’ life is 3 to 6 months. Beetles which stay for wintering live up to 8 months; the places for the wintering are piles of grain, underground galleries, burrows of rodents and other secluded places in non-heated buildings (Ganiev et al., 2009). The rice weevil is more fertile than the wheat weevil, and lays up to 500 eggs. The way of laying eggs and further development are the same as of the granary weevil. The rice weevil is a thermophile, and cold temperatures are lethal for it. In southern areas, over a year it has up to 4, and in the northern – up to 2 generations. Optimum temperature for the development of larvae is 28–30 °С, and grain moisture is 18 %. In these conditions, the development cycle lasts 23–25 days. At 21–25 °С, the development of one generation lasts about 40 days, whereas at 14–18 °С it slows to 3.5–7.0 months. At the environmental temperature below 13 °С and moisture of grain (wheat) below 10%, the rice weevil does not develop. Rice weevils harm grain of rice, wheat, rye, maize, barley, flour, wild bean, seeds of cannabis, pearl barley and dry flour products, seeds of panic grass, essential oil-bearing plants and bean plants. Compared to the wheat weevil, the rice wee- 76 Chapter 2. Coleoptera pests of stored food supplies and field crops vil causes much more damage. Due to the development of the rice weevil, cereal grains lose 35 to 75 % of their weight. The breeding grounds are plant remains with grain of different cereals which ac- cumulate over years on threshing floors. Annual accumulation of unsuitable grain wastes allows the beetles to survive in them over a long time. In cases when the food supplies in the sources are exhausted before the harvest or when old threshing floors are taken out of use, the weevils leave them. At the same time, they make mass flights before the grain harvest (Eremenko et al., 1967). There have been occurrences of infestation of grain in fields at distances of up 1.5 km from stores with infested grain (Sokolov, 2004). Cosmopolite. Introduced in all countries around the world. Distributed in Europe, the Mediterranean, the Oriental region, Oceania, the European part of Russia. Can be found throughout Ukraine in stores of grain and products of its processing. In field conditions, it can be found in the tropical belt. The beetles feed on wheat, rye, barley, oat, sorghum, cannabis seeds, millet, corn, rice, flour, bran, rusks, cookies, macaroni, dried apples and other products; however, the larvae do not develop in flour, bread, cannabis seed, millet, bran, and dried apples. Sitophilus zeamais Motschulsky, 1855. Body size – 2.5–5.0 mm. The beetle looks similar to S. oryzae, but is larger. The upper part of the body is semi-mat with slight gloss, light spots on the cuticle at the base of the elytra reach the shoulder angles (Sokolov, 2004). In- dividuals of typical coloration have red-brown spots on the elytra distinctly defined, brighter compared to wheat and rice weevils; the spots in the front part of the elytra usually extend to the shoulder tubercles. The pronotum has coarse dense rounded punctures, some of the punctures are sharply impressed, with bright gloss; the areas between them are narrow, carinate-sharpened, glossy (Shamonin, 1986). The beetle has two pair of wings and is able to fly (Eremenko et al., 1967). The development of Sitophilus zeamais or maize weevil is similar to the development of S. oryzae. In resistance to cold, the maize weevil has an intermediary position behind S. granaries, but superior to S. oryzae. For example, at 5 °С, the beetles of wheat/granary weevil lived 30 days, while maize beetles – 23, and V. I. Rusynov, V. O. Martynov, T. M. Kolombar 77 rice weevil – 4 days (Sokolov, 2004). Distributed in moderately warm countries of America, Europe, Asia and Africa. Causes the same damage as other weevils. Harms grain of wheat, rye, oat, barley, rice, corn, buckwheat, pearled barley and other flour products, seeds of panic grass, essential oil-bearing plants and bean crops. Grain damaged by the weevil becomes unsuitable for food (Ganiev et al., 2009).

Family Meloidae

Subfamily Meloinae Meloe (Lampromeloe) variegatus Donovan, 1793. Body size – 11–42 mm. The body is large and strong, the elytra are short, at the bases one overlaps the other, further diverging. The abdomen is very long, swollen; extends beyond the apex of elytra, sometimes among males, and always among females. The upper part is coarsely wrinkled, bronze, green or red-violet, the abdominal terga have copper-red transversal stripes. The head behind the eyes often has a longitudinal furrow or thin keel. The antennae are not thickened in the middle. The pronotum is usually transversal. The posterior coxae are no longer than their width. Distributed in South and Central Europe, North Africa, the European part of Russia, Siberia and the Near East (Luckmann and Scharf, 2004). Can be found throughout Ukraine. The beetles sometimes gnaw leaves of sugar beet, cabbage, rye, wheat. Larvae can cause damage to beekeeping, and cause parasitic diseases of bees. Meloe (Meloe) proscarabaeus Linnaeus, 1758. Body size – 13– 22 mm. The antennae of males are significantly thickened in the middle, the antennae of females are notably thickened near the base and become narrower towards the top. The width of the pronotum is no greater or almost greater than the length. The posterior coxae are longer than their width. The head and the pronotum bear coarse dense punctures. The elytra have coarse skin-like wrinkles, and are quite large in the shoulders. Black with bluish shimmer, more rarely dark blue. The antennae of males are curved in the middle (Di Gi- ulio et al., 2014). Distributed in Europe, the Mediterranean, the Eu- 78 Chapter 2. Coleoptera pests of stored food supplies and field crops ropean part of Russia, the Caucasus, Siberia, Central Asia and the Near East. Can be found throughout Ukraine in various habitats. The beetles feed on leaves of Ranunculaceae, Fumarioideae, Astera- ceae, spurge (Euphórbia), Compositae, Umbrelliferae, sometimes gnaw leaves of beet, potato, clover, stems of seedlings of cereals, mustard, and sunflower. Meloe (Meloe) violaceus Marsham, 1802. Body size – 10–32 mm. The head and the pronotum are slightly and sparsely punctured. The elytra are very narrow at the shoulders, with longitudinal wrinkles. Blue or violet, very rarely black (Lückmann and Assmann, 2006). Distributed in Europe, the Mediterranean, North Africa, the Euro- pean part of Russia and the Near East. Can be found throughout Ukraine in forest edges, forest glades, gardens, meadows, swampy areas, can feed on the leaves of potato, clover, sprouts of wheat, maize. Meloe (Eurymeloe) scabriusculus Brandt & Erichson, 1832. Body size – 9–28 mm. The head and the pronotum bear very thin hairs seen from the side against the light, densely and significantly wrinkled- punctured. The antennae are thinner and longer, not thickened near the top. Black-blue or blue, slightly glossy. The eyes are small and flat. The elytra are densely and finely pebbly. Distributed in Alba- nia, Austria, Bulgaria, Croatia, the Czech Republic, France, Germa- ny, Greece, Hungary, Macedonia, the Near East, Poland, Romania, Slovakia, Slovenia, Switzerland, Moldova, the Caucasus and Cen- tral Asia (Luckmann and Scharf, 2004). Can be found throughout Ukraine. Has been observed to damage the sprouts of wheat. Mylabris (Eumylabris) crocata Pallas, 1781. Body size – 7–16 mm. The antennae thicken towards the top. The mandibles are asymmetric. The claws are usually not serrated. The elytra usually have membranous sashes or rows of spots, the rest of the body is usually black or metallic, densely haired. The arrangement of the spots on the elytra: 2, 2, 2. The larvae develop in the egg cap- sules of locusts, more rarely – in nests of solitary bees. The larvae feed on flowers, more rarely on leaves, can damage leaves of soybean, lucerne and other plants. Distributed in Europe, Iran, Syria, European part of Russia, Siberia, Kazakhstan and Central V. I. Rusynov, V. O. Martynov, T. M. Kolombar 79

Asia (Hayat et al., 2002). In Ukraine, can be found in Kirovohrad, Kherson Oblasts and Crimea. Mylabris (Eumylabris) fabricii Soumacov, 1924. Body size – 9–20 mm. The pronotum is conically narrowed towards the top, has a depression or longitudinal furrow in the middle, its disc is often sculptured. The pattern on the elytra is composed of rounded spots. The arrangement of the spots on the elytra: 2, 2, 1. Distributed in Europe, Iran, the south of the European part of Russia, and in the Near East. In Ukraine, can be found in the steppe zone and Crimea. The beetles most often concentrate on wild-growing Compositae and Cruciferae, but can also harm crops by gnawing flowers and young leaves of cabbage, radish (Raphanus raphanistrum subsp. sativus and Raphanus sativus var. radicula), mustard, lucerne, cereals (Zhu and Higgins, 1994). Mylabris (Mylabris) quadripunctata Linnaeus, 1767. Body size – 11–16 mm. The elytra are red or orange with black apices and two pairs of spots - in the first quarter and in the middle section; sometimes the posterior pair fuses. Distributed in South Eu- rope, Turkey and Western Asia, European part of Russia, Central Asia (Luckmann and Scharf, 2004). In Ukraine, can be found in the steppe and forest-steppe zones, in highlands of Crimea, and around Kyiv and Lviv. Habitat generalist xerophile. The beetles harm the ears of rye, flowers of poppies, safflower (Carthamus tinctorius), flowers and leaves of mustard, rape, flowers and seeds of cabbage, dill, pumpkin, melon, soybean, lucerne, sainfoin, sunflower. Mylabris (Micrabris) sibirica Fischer von Waldheim, 1823. Body size – 7–11 mm.The pronotum is non-uniformly pebbled, with a low smooth keel. The pattern is similar to that of the previous beetle, but the apical fringe is nearly always present (Bey-Bienko, 1965). Distributed in North-West Mongolia, the European part of Russia, Siberia, Kazakhstan, Kyrgyzstan. Can be found throughout Ukraine, except Polesia. Meadow mesophile. Can consume anthers of wheat, barley, sugar beet, flowers of lucerne, clover, vetches, Compositae and Cruciferae. 80 Chapter 2. Coleoptera pests of stored food supplies and field crops

Mylabris (Mylabris) variabilis Pallas, 1781. Body size – 10–17 mm. The pronotum is uniformly and largely punctured. Dis- tributed in South Europe, Asia Minor, North Africa, the European part of Russia, the Caucasus, Transcaucasia, Kazakhstan, Central Asia. In Ukraine, can be found in the steppe and forest-steppe zones, Zakarpattia Oblast and Crimea. The beetles are habitat generalist xerophiles, can consume the flowers and young leaves of garden crops of Cruciferae, Asteraceae, and Poaceae.

Family Tenebrionidae

Subfamily Alleculinae Omophlus flavipennis Kuster, 1850. Body size – 7.5–12.0 mm. Black, the elytra are yellow-brown or red-yellow. The lower part of the body and the femora are covered with homogenous light-grey hairs. The internal claw of the forelegs of the male has a tooth; the fifth segment of the forelegs of the male is enlarged; the main segments of the antennae, palpi, protibiae and feet are yellow. Distribut- ed in Italy, Romania, Hungary, Dalmatia, Turkey and Transcaucasia (Kiliç and Yildirim, 2009). In Ukraine, can be found in the south of the steppe zone and in Crimea. The life cycle and extent of damage are the same as for O. proteus. Omophlus lividipes Mulsant, 1856. Body size – 6.5–9.5 mm. The lower part of the body and femora bear dense black appressed and more rarely protuding hairs. The pronotum has fine yellowish appressed and more rarely partly appressed hairs. The antennae of males equal 3/4 of the body length, the forelegs are notably thickened. The main segments of the antennae, tibiae and the feet are yellow-brown. The lateral margins of the pronotum are rounded, narrow in the front, largely extending outward. Distributed in Austria, Germany, Italy, Poland, the European part of Russia, Ciscaucasia, Kazakhstan (Nakládal et al, 2017). In Ukraine, is more often found in the Central and Left-bank forest-steppe. Is distinct from the other species of this genus in having dark-yellow coloration of the main part of the antennae, tibiae, V. I. Rusynov, V. O. Martynov, T. M. Kolombar 81 and feet. The life cycle and extent of damage are the same as for O. proteus and O. flavipennis. Omophlus proteus Kirsch, 1869. Body size – 7.0–14.5 mm. The elytra have obvious puncture striae, with glossy smooth or slightly wrinkled intervals; the punctures in the furrows are larger compared to the intervals. The main part of males’ elytra has well notable protruding hairs, the rest of the elytra of males and females is bare. Dis- tributed in Europe, Turkey, the European part of Russia. In Ukraine, can be abundant in the belt of forbs and Volga fescue (Festuca valesiaca) – feather grass (Stripa) steppes, and in Crimea (Ghilarov, 1937). The beetles concentrate during the day on the flowers of different plants, often in large numbers, feed on pollen and different organs of flowers, preferring the Rosales family, but can also eat the anthers of Poaceae, Cruciferae and other plants, often gnawing the petals and pistils. Young larvae are saprophages. Older larvae also prefer dead plant remains, but can also damage living plants, mainly the seedlings of different crops. Can be found in numbers ranging from several dozen to 200 individuals/m2, however usually causing no significant damage to crops.

Subfamily Diaperinae Gnathocerus cornutus Fabricius, 1798. Body size – 3.2–4.5 mm. The body is parallel-sided, bare. The pronotum in the front is of the same width as the elytra. The elytra have very thin puncture striae. Rusty-red, insignificantly glossy. Cosmopolite. Distributed in Aus- tria, Belgium, Great Britain, Croatia, the Czech Republic, Denmark, Estonia, Finland, France, Germany, Hungary, Italy, Latvia, Lithuania, Norway, Poland, Sardinia, Sicily, Slovakia, Sweden, central and south belt of Russia (Tsuda and Yoshida, 1985). Can be found throughout Ukraine. Found in flour, but it seems that it feeds not only on flour, but rather consumes other storage pests – larvae of flour beetles.

Subfamily Palorinae Palorus depressus Fabricius, 1790. Body size – 3 mm. The body is elongated, brown-red. The head has deep transversal impressions 82 Chapter 2. Coleoptera pests of stored food supplies and field crops between the clypeus and the forehead. The pronotum is insignificantly transversal, widest behind the front margin, from where it narrows almost directly towards the base. The elytra bear thin puncture striae. The largest impression is the one which stretches along the front edge of the head and reaches the front edge of the eye. The side edge of the cheek reaches the front edge of the eye, and contin- ues further (to the middle of the upper part of the eye) only as a very thin, hardly notable, keel; practically the entire upper part of the eye is seen from the top. The upper edge of the head above the antenna is almost straight. The clypeus bears rare, significantly erased punctures. The puncture row in the third interstria of the elytra is sometimes doubled (Bey-Bienko, 1965). Distributed in the middle and south parts of the Eurasia, central and south areas of the Euro- pean part of Russia (Shavrin et al., 2015). Can be found throughout Ukraine. Develops in rotten wood, under the bark of deciduous and coniferous species; mostly predatory. Can be found also in storage facilities on grain, grain products, in flour and flour products. Palorus ratzeburgi Wissmann, 1848. Body size – 2.8–3.0 mm. The clypeus bears small but distinctive punctures. Puncture stria in the third interstria of the elytra is not doubled. Distributed in Austria, Belgium, Great Britain, Croatia, the Czech Republic, Denmark, France, Germany, Greece, Hungary, Italy, Latvia, Norway, Poland, Sardinia, Sicily, Slovakia, Sweden, in the south of the European part of Russia (Cooper and Fraenkel, 1952). In Ukraine, it can be found in the southern steppe zone in stores of grain and grain products. Palorus subdepressus Wollaston, 1864. Body size – 3 mm. The impression along the front and lateral edge of the head reaches the posterior edge of the eye. The lateral edge of the cheek also reaches the posterior edge of the eye, and therefore the upper edge of the eye cannot be seen from above. The edge of the head above the antennae is gradually rounded. The furrow between the clypeus and the cheek is smoothened. Distributed in Great Britain, Croatia, Denmark, Finland, France, Germany, Greece, Hungary, Italy, Poland, Sardinia, Sicily, Slavakia, Sweden, the south of the European part of Russia (Doumbia et al., 2014). In Ukraine, it can be found in the steppe V. I. Rusynov, V. O. Martynov, T. M. Kolombar 83 zone and in the southern forest –steppe zone. Harms grain and grain products. Breeds in stores. Subfamily Pimeliinae Asida lutosa Solier, 1836. Body size – 9–12 mm. Black, covered in appressed reddish hairs, between which particles of soil usually stick. The pronotum is densely punctured, with impression in front of the scutellum. Each elytron on the sides bears a sharp keel and four insignificant intermittent wavy ribs. Distributed in Bulgaria, Romania, and the Caucasus. In Ukraine can be found in the steppe zone (Ghilarov, 1937). The beetles and larvae prefer virgin lands and set-aside lands, where it occurs in large numbers. On cultivated lands, the beetle is found rarely or singularly, mostly in tilled areas, sometimes damages cultivated plants similarly to T.nomas. Tentyria nomas Pallas, 1781. Body size – 12–15 mm. Black, glossy. The elytra are smooth or with thin longitudinal furrows, sometimes with transversal wrinkles. The epipleura of the pronotum are covered in coarse and deep longitudinal wrinkle-furrows. Dis- tributed in the European part of Russia, Ciscaucasia, Kazakhstan, Central Asia. In Ukraine, the beetle can be found in the south forest- steppe and in the steppe zone. Prefers virgin lands and abandoned lands, where the larvae develop. The beetles crawl into ploughed lands, where they cause damage to agricultural plants, mainly on tilled ground , insignificant harm similarly to O. sabulosum. The larvae on ploughed soils are observed rarely or in single specimens and can harm seedlings and sprouts (Ghilarov, 1937). Pimelia subglobosa Pallas, 1781. Body size – 14.0–18.5 mm. Black, the elytra are oviform or ball-shaped. The protibiae have a tooth on the top of the external edge. The apical tooth of the protibiae is sharpened. The middle and hind legs are not doubled. The elytra are ball-shaped, with large smooth tubercles, have small erasing hairs between the tubercles. The side rib of the elytra is significantly swollen. Distributed in Bulgaria, Turkey, Greece, Macedonia, Roma- nia, the south of the European part of Russia (Leo and Fattorini, 2000). In Ukraine, can be found in the steppe zone. The beetles harm 84 Chapter 2. Coleoptera pests of stored food supplies and field crops agricultural crops in a similar manner to owlet moths, and the larvae can damage seeds and the sprouts similarly to the larvae of the Blaps genus. The beetles and larvae prefer virgin lands, and are not observed in large numbers in fields with crops. Subfamily Tenebrioninae Alphitobius diaperinus Panzer, 1797. Body size – 5.5–6.5 mm. Oblong-oval, bare. The elytra have puncture rows, deeper in the upper part. The protibiae are enlarged towards the top. The pronotum is widest in the base or in the middle, but in this case it slightly narrows towards the base. The puncture rows on the apex of the elytra turn into furrows. Black or brown. Distributed in Austria, Belgium, Crete , Croatia, Cyprus, the Czech Repub- lic, Denmark, Estonia, Finland, France, Germany, Great Britain, Greece, Hungary, Italy, , Latvia, Lithuania, Malta, Norway, Poland, Sardinia, Sicily, Slovakia, Sweden (Salin et al., 2000; Chernaki-Lef- fer et al., 2007). Found in storage premises throughout Ukraine. Alphitobius laevigatus Fabricius, 1781. Body size – 4.5–5.0 mm. The pronotum has significantly rounded sides, narrowing slightly less towards the base rather than to the front. The puncture rows on the top of the elytra do not change into furrows (Fox and Bayona, 1968). Distributed in Belgium, Great Britain, Cyprus, the Czech Re- public, Denmark, Estonia, Finland, France, Germany, Greece, Italy, Latvia, Madeira, Malta, Norway, Poland, Sicily, Slovakia, Sweden, but occurs rarely (Maitip et al., 2016). Can be found throughout Ukraine in stored grain and flour. Blaps halophila Fischer de Waldheim, 1820. Body size – 17–23 mm. The body is usually large, bare on the upper part. The elytra have a long tail-like projection in the top. The antennae are shorter, their apices do not reach the basis of the pronotum, the first segment of hind legs is notably asymmetric. The pronotum is swollen, insignificantly transversal (width is 1.1 times greater than the length) (Allsopp, 1980). Distributed in Austria, Bulgaria, Croa- tia, the Czech Republic, Hungary, Poland, Romania, Slovakia, south of the European part of Russia, Siberia, Kazakhstan, Tajikistan, V. I. Rusynov, V. O. Martynov, T. M. Kolombar 85

Uzbekistan (Chigray et al., 2016). In Ukraine, can be found in the steppe zone and Crimea. The beetles are active in the morning and evening hours, during the day they hide in shelter, crawl into burrows of rodents, basements, storage premises and other dark places, under sheds and different structures, including houses. In the farm fields, the beetles similarly to B. lethifera feed on withered plants, cut during mechanical processing the rows between tilled crops. In barns and storage places, the beetles gnaw seedlings and sometimes damage potato tubers, and also feed on different grain remains (barley, wheat, maize and other food supplies). The development of larvae and the damage they cause is similar to B. lethifera. The greatest damage is caused by wintering larvae to sown seeds of maize and other cereals, sugar beet, sunflower, Cucurbitaceae, and also the seedlings of agricultural plants. In places of concentration (breeding grounds), larvae of such species as B. lethifera can cause serious damage to crop fields. Blaps lethifera Marsham, 1802. Body size – 20–27 mm. The protibiae are narowed at the base, but have no groove]. The pronotum bears small, moderately dense punctures, the distance between the punctures on the disk is smaller than the punctures. The apical slope of the elytra is very steep. The side fringe of the pronotum is thin, not thickened in front of the basis. The pronotum is rarely heart-shaped. The elytra are swollen. Distributed in Albania, Aus- tria, Belgium, Great Britain, Corsica, Croatia, the Czech Republic, Denmark, Estonia, Finland, France, Germany, Greece, Hungary, It- aly, Crete, Madeira, Norway, Poland, Romania, Sardinia, Sicily, Slo- vakia, Sweden, Asia Minor, the south of the European part of Russia, Siberia, Kazakhstan, Uzbekistan, the Caucasus. In Uktaine and the south forest-steppe, in the steppe zone, in arable lands and virgin lands (Dekhtiarev, 1928). The larvae cause damage in soil. They inhabit illuminated areas of fields, prefer tilled crops and garden crops, more rarely occur in wheat and barley. Active in the morning and evening hours, and during the day, they hide in shelter and crawl into burrows of rodents, storage places and basements. The beetles feed on the seedlings of different weeds (saltbush (Atriplex), Sal- 86 Chapter 2. Coleoptera pests of stored food supplies and field crops sola ruthenica Iljin.and others), in beet fields, they prefer feeding on withered plants which remain after weed control and thinning (Allsopp, 1980). Blaps tibialis Reiche, 1857. Body size – 20–24 mm. The apical projections of the elytra of males are short (up to 2.0) and in females undeveloped. The I sternum of the abdomen of males is positioned between the coxa and the tubercule. The protibiae in the base on the internal margin has a notable incisure (less significant in females than in males). The pronotum bears dense, partly fused punctures, the distance between the punctures is smaller than the punctures themselves. Distributed in Bulgaria, the European part of Turkey, Greece, Crete, Romania, the European part of Russia (Soldati et al., 2017). In Ukraine, occurs in the southern steppe zone. Observed in floodplain steppe areas along with B. lethifera, but always as single specimens. The way of life and damage caused by these beetles and their larvae are similar to B. lethifera. Gonocephalum granulatum Fabricius, 1791. Body size – 6.5–8.0 mm. The protibiae are only slightly wider than the middle, the apical tooth of the tibia does not reach farther than the third segment of the leg. The third segment of the antennae is 2.0–2.5 times longer than the second. The dorsal part is almost entirely bare, with hardly noticeable setae. The uneven intervals of the elytra usually protrude further than the even intervals. Distributed in Albania, Austria, Bulgaria, Corsica, Croatia, the Czech Republic, France, Greece, Hungary, Italy, Macedonia, Romania, Sardinia, Sicily, Slovakia, Spain, Asia Minor, the south of the European part of Rus- sia, the Caucasus, Kazakhstan, in mountains of Central Asia (Bunal- ski et al., 2014). In Ukraine, can be found in the steppe zone, most abundant in south subzones of southern chernozems and dark kas- tanozems. The damage to the plants caused by the beetles and larvae of this species is of the same type as damage caused by O. sabulosum. Opatrum sabulosum Linnaeus, 1761. Body size – 6.5–10.0 mm. The base of the pronotum is of the same width as the base of the elytra, or slightly wider. Indistinctive intervals of the elytra are raised. The pronotum is uniformly granular, without smooth elevations. V. I. Rusynov, V. O. Martynov, T. M. Kolombar 87

The cant of the pronotum is moderately raised. Distributed in the European part of Russia, Siberia, the Caucasus, Kazakhstan, Asia Minor (Medvedev, 1968). Can be found throughout Ukraine. The larvae feed on rotting plant remains, practically do not harm live plants or even if present in large numbers. The beetles are polyphagous and damage seeds of various crops, but are most harmful to the seedlings of tilled crops and seedlings of garden crops in spring and early summer. Over this period, several dozens to several hundreds of beetles can concentrate on one square meter. Especially noticeable is the damage it causes to sunflower, maize, tomato, cucumber, soybean, wild bean, onion. The beetles prefer feeding on plants which are beginning to wither, and therefore are especially harmful for freshly sown seedlings of garden crops and tobacco. Beetles eat the leaf laminae of grasses, and cotyledons of sunflower and other crops (Brygadyrenko and Nazimov, 2015). Episodes have been observed of the beetles damaging swollen and sprouting seeds of barley, wheat, oats, Cucurbitaceae, and other plants. The most significant damage is caused by the beetles in late April to mid May. By early June, the intensity of damages diminishes and by mid-June it has practically stopped. Tenebrio molitor Linnaeus, 1758. Body size – 12–16 mm. The head is small, rounded. Black-brown, with fatty gloss. The last segment of the antennae is not transversal, semi-rounded. The head and the pronotum are densely punctured. The pronotum at the base has no swollen stretched part, only a deep transversal furrow, the ends of which are orientated to the front, the posterior angles of the pronotum are sharpened or straight. The elytra have obvious furrows, the punctures in the furrows are not always distinct. Cosmopolite, considered native to the Mediterranean. Distributed throughout Ukraine. Storage pest. Beetles and larvae damage products of grain milling (flour, especially with heighted moisture or fusty groatы and brans), and also grain, rusks, dairy products (macaroni, vermicelli), etc (Martynov and Brygadyrenko, 2017). In the grain, the larvae first consume the embryo, and then the endosperm. The damage caused by the beetles consists not only in their consumption of part of the 88 Chapter 2. Coleoptera pests of stored food supplies and field crops product, but mainly in their contaminating the products with ex- crement and larvae skin. Damaged products lose their capacity for long-term maintenance, obtain an unpleasant smell, and the damaged grain loses ability to sprout. Inhabits mills, grain stores, confectionery and macaroni factories, bakeries, any places where the sanitary requirements are not fulfilled, and also humid unventilated places, where food spills out of its packaging (Sokolov, 2004). Tenebrio obscurus Fabricius, 1792. Body size – 14.0–18.5 mm. The throat has no lateral spikes. Black, completely mat. Last segment of the antennae is transversal. The dorsal part is covered in numerous fused punctures. The pronotum has a cylindrical section at the base. The elytra have barely noticeable puncture rows, the intervals have large scattered flat grains. Distributed throughout the forest and forest- steppe zones (Cotton, 1927). In Ukraine, can be found in Polesia and the forest-steppe. Sometimes is found in stores, but usually in natural conditions. Larvae develop in rotten wood and more rarely in forest litter, singularly occur in tilled fields. Is not numerous. Tribolium castaneum Herbst, 1797. Body length – 3.5–5.0 mm. The beetle is red-brown to rusty-red in color, almost mat; the pronotum is uniformly rounded on the sides, not grooved; it is widest in the middle; the thorax scutellum bears numerous small punctures; the elytra are puncture-furrowed; the intervals bear very small punctures (Sokolov, 2004). Tribolium castaneum or the red flour beetle is very similar to the mealworm beetle (Tenebrio molitor), and is distinguished from it by glossy red-brown color and much smaller size (Eremenko et al., 1967). Also, the red flour beetle look similar to the confused flour beetle (Tribolium confusum), distinguished from it by the following features: three terminal segments of the antennae are sharply thickened; in the lower part of the head the distance between the eyes equals the width of the eye; the wings are well developed, beetles fly well, especially at night (Dashevskiy and Zakladnoy, 1978). The red flour beetle is more thermophilic than the confused flour beetle. Prefers bread and feed grain with damaged grains and seeds of weed, which the beetles and larvae feed on. By this feature, the beetle is distinct from confused flour beetle which occurs more often in flour V. I. Rusynov, V. O. Martynov, T. M. Kolombar 89 sweepings and flour dust. Larvae are white at first, but become yellow in the terminal stage, are 3–7 mm in length, with two triangular hook- like projections on the 9th abdominal segment. The pupa is yellow- white, looks similar to the pupa of the mealworm beetle, but its length is no more than 3.5 mm. The female pupa has two small tubercles at the end of the abdomen, and the male pupa has one small prominence (Sokolov, 2004). In favorable conditions, the females are able to lay 300–350 eggs on average over their life, and 1000 eggs maximum. Eggs are hard to find due to the fact that they are covered with sticky liquid which quickly becomes covered in particles of flour and flour dust. The development of eggs at 25 °С lasts 6–7 days. Larvae undergo 5–12 moults, transform into pupae, and then into beetles. The larvae develop best in flour and bran. In cases of mass contamination, the flour becomes lumpy. The flour contaminated with larvae obtains a dirty color, unpleasant smell and taste and is unfit for human consumption (Bey-Bienko, 1966). Depending on the temperature, the larvae develop in 20–100 days. At the temperature of 27 °С, the pupa phase lasts 10–12 days. The lower temperature threshold for the development of red flour beetle is 15.2 °С. At the temperature of 25 °С, the development cycle lasts 47 days. At 30 °С, in flour, the development of eggs, larvae and pupae lasted 4, 19 and 5 days respectively. As with confused flour beetle, it develops throughout the year in heated premises. In unheated premises, the beetles winter. The beetles have a long life span of up to two years and occasionally longer (Eremenko et al., 1967). The beetles are extremely tolerant of low content of moisture.In ground products, they develop at moisture content of around 1 %. The abundance of any given population of red flour beetle is limited by the fact of cannibalism, when beetles and larvae eat eggs and pupae of their own species. The cannibalism increases as the density of the insects’ population increases. Is distributed with contaminated products, containers and storage equipment; within the territory of one enterprise actively distributes itself during night flights. Cosmopolite. Distributed in the southern regions of European part of Russia, Central Asia. Distrib- uted throughout Ukraine. Damages groats, compound feed, bran and 90 Chapter 2. Coleoptera pests of stored food supplies and field crops other ground grain products. Prefers bread and feed grain with pul- verised grain and seeds of weeds, which the beetles and larvae feed on (Sokolov, 2004). Often found together with confused flour beetle in batches of grain, flour and groats. Tribolium confusum Jacquelin du Val, 1863. Body size – 3.5–5.0 mm. Confused flour beetle looks very similar to the mealworm beetle, differing from it in glossy red-brown color and significantly smaller size. Also, the confused flour beetle looks similar to the red flour beetle, differing from it in the following parameters: the antennae gradually thicken towards the top; in the lower part of the head, the distance between the eyes is 3 times wider than the width of the eye; the wings are undeveloped, beetles are flightless (Dashevskiy and Zakladnoy, 1978). The prothorax is rectangular in the upper part. The rounded end of the abdomen of male bears hairs and is bare in females (Eremenko et al., 1967). The eggs are white, of short oval shape, 0.7 mm long. The larva is flattened, white at first, but becomes yellow in the final stage, body length is 3–7 mm, has three hook-like projections on the 9th abdominal segment. Adult larva is yellowish above, lighter below, covered in sparse long hairs (Shorohov and Shorohov, 1936). The pupa is yellow-white, looks similar to the pupa of the mealworm beetle, but its length is no greater than 3.5 mm. The female pupa has two small tubercles at the end of the abdomen, and the male pupa of has a small elevation (Sokolov, 2004). Over unfavorable conditions, females of confused flour beetles can lay on average 300–350 eggs over their lifespan, and up to 1,000 eggs maximum. Eggs are hard to find, for they are covered in sticky liquid which collects particles of flour and dust. The development of the eggs lasts 6–7 days at 25 °С. The development of the embryo in the egg stops at temperatures below 10 °С, the development cycle of the confused flour beetle is completed in 56 days on average. Larvae undergo 5–12 moults, transform into pupae and then into beetles. The larvae develop best in flour and bran (Zakladnoy et al., 2003). In cases of massive contamination, the flour becomes lumpy. The flour contaminated with larvae obtains a dirty color, unpleasant smell and taste, and becomes unfit for con- V. I. Rusynov, V. O. Martynov, T. M. Kolombar 91 sumption. Depending on the temperature, larvae develop over 20– –100 days. At the temperature of 27 °С, the pupa phase lasts around 10–12 days. The beetles live, eat and breed for up to three years. The lower temperature threshold of the development of the confused flour beetle is 14.8 °С. In favorable temperatures of 23–25 °С, the development from egg into an adult beetle takes 35–45 days. At the temperature of 25 °С, the development of the confused flour beetle is completed on average in 56 days. At 30 °С, in flour, the development of eggs, larvae and pupae of the confused flour beetle is completed in 6,25 and 6 days respectively. In heated premises, confused flour beetles can produce up to 4 generations over a year, in unheated – 1–3. The abundance of any given population of the beetles is limited due to the cannibalism, when the beetles and larvae consume eggs and pupae of their own species. Cannibalism increases as the insects’ population density increases. The insects are highly tolerant to low content of moisture. In milled products, they can develop at moisture content around 1 % (Zakladnoy et al., 2003). Confused flour beetle is a thermophilic species. Even at 0 °С, it lives only 2–3 days. Typical inhabitant of heated premises. Cosmopolite. Distributed in the south of the European part of Russia, Central Asia. Quarantine object in Mongolia, Slovakia and Hungary. Distributed throughout Ukraine. The damage caused by the beetles and larvae of confused flour beetle consists in their consuming and contaminating various products, especially flour, to which they are particularly attracted . Found in mills, feed compound factories, bread factories, bakeries, confectionery and macaroni factories, breweries and food supplies stores. Feeds on flour, groat and bran, rarely damages wheat, dried vegetables and fruits. Absolutely does not feed on grain legumes and solid grains of crops with scarious bracts - oats, barley and rice (Sokolov, 2004). Con- fused flour beetle also feeds on grain, but only if the grain is damaged; it cannot consume healthy, solid grains, and dies. It should be mentioned that it usually damages the upper layer of heaped products (Shorohov and Shorohov, 1936). The beetles have scent glands on the thorax and abdominal segments, which secrete fluid 92 Chapter 2. Coleoptera pests of stored food supplies and field crops with a sharp unpleasant scent, which contains quinones. Especially significant damage is caused in mills, where it develops throughout the year, nests inside different mechanisms which are hard to clean (Eremenko et al., 1967). Tribolium destructor Uyttenboogart, 1934. Body size – 5.1–5.5 mm. Black. The angle of the cheek near the front edge of the eye is rounded. The front edge of the pronotum has rounded significantly prominent angles. Cosmopolite (Brown, 1950). Can be found throughout Ukraine. Damages various grain products, allspice, glue, knitted and nylon goods. Inhabits only heated premises, including apartments, where produces 3–4 generations during the year. Way of life similar to T.confusum, but is less harmful. Less tolerant to cold than T.castaneum. Tribolium madens Charpentier, 1825. Body size – 4–5 mm. Three terminal segments of the antennae have a clearly distinct capitulum, the head has no keel at the internal edge of the eye. Dark-brown to black. The width of the pronotum is 1.4 times greater than the length. Only the first interstria of the elytra has no keel, the second interstria at the base has a keel. Distrib- uted in Central and Southern Europe, European part of Russia (Halstead, 1969). Distributed throughout Ukraine. Similarly to T. confusum, together with which it is usually found, causes damage to the same products, leads a similar way of life to this species, but causes less damage.

Family Dascillidae

Subfamily Dascillinae Dascillus cervinus Linnaeus, 1758. Body size – 10–12 mm. Is almost parallel-sided, black. Males bear dense grey hair; the elytra, antennae and legs of females are yellow-brown, the top has dense yellow hairs (Baker, 1981). Distributed in the European part of Rus- sia, except the tundra and steppe zone. In Ukraine, is distributed in forest and forest-steppe zones. The larvae harm roots of oats, grasses and trees. V. I. Rusynov, V. O. Martynov, T. M. Kolombar 93 Family Elateridae

Adrastus rachifer Fourcroy, 1785. Body size – 3.0–3.5 mm. Black or dark-brown, the antennae, legs, the front edge of the pronotum and its posterior edges and the elytra, except the stripe along the elytral suture and sometimes the sides are yellow. Distributed in the south of the European part of Russia, the Caucasus (Kabalak and öZbek, 2018). In Ukraine, can be found in forest-steppe zone; through the river valleys it travels to the steppe zone, including steppe Crimea. In leached chernozems in lowlands, forms concentrations of larvae of up to 30 larvae per 1 m2, including in crop fields. Agriotes brevis Candeze, 1863. Body size – 6.5–9.0 mm. The width of the pronotum is slightly greater than the length, the elytra are not enlarged in the posterior third part. Black or black-brown, the antennae, legs and a wide stripe, which sometimes disappears along the edge of the elytra, are red-brown. Distributed in Asia Minor (Tóth etal., 2002). In Ukraine, can be found in Zakarpattia Oblast. In the lowland part of Zakarpattia Oblast, causes damage together with A. sputator, from which it is distinguished by the transversal pronotum and glossy surface of the body. Life pattern and harmfulness of larvae are similar to A. sputator. Agriotes gurgistanus Faldermann, 1835. Body size – 10–14 mm. Swollen, the pronotum enlarges towards the posterior angles. The greatest width of the middle segments of the antennae is 1.5 times less than their length. Black-brown, sometimes of chestnut color, the antennae, legs, lower part, sometimes only the edges of the segments, are lighter; the upper part is covered in silky grey indumentum. Distributed in Bulgaria, Romania, Asia Minor, the south of the European part of Russia, the Caucasus (Kudryavtsev et al., 1993). In Ukraine, is most abundant in south forest-steppe, in north and middle belt of the steppe zone, some concentrations were found in the southern steppes. The formation of concentrations of larvae of this species is not related to grasses. Damage is caused by larvae of all ages, but especially harmful are larvae of final stages in plantations of maize and garden crops. 94 Chapter 2. Coleoptera pests of stored food supplies and field crops

Agriotes sputator Linnaeus, 1758. Body size – 6.0–8.5 mm. The prothorax is punctured less coarsely and densely compared to its epipleura. Areas between the puncture of prothorax are at least twice as wide as the diameter of the punctures. Width of the pronotum equals its length. The elytra are slightly enlarged in the posterior third part. Dark- or light-brown in grey indumentum, the pronotum is usually slightly darker than the elytra, the antennae and legs are lighter. Distributed in Asia Minor, North Africa, the European part of Russia, the Caucasus, Siberia, Kazakhstan (Benefer et al., 2012). Distributed throughout Ukraine, the zone of its highest abundance includes the entire forest-steppe and the northern strip of the steppe zone. One of the most abundant species of pests in tilled lands of the middle belt (Čačija et al., 2017). The highest number of its larvae was recorded in sod loamy, sod-podzolized, manure-carbonate, grey forest soils, degraded and thick low-humus chernozems, where their number often reaches several dozen per 1 2m , sometimes even reaches 100 specimens and higher. Damage is caused by larvae of all ages, but the most harmful are larvae of final stages. The formation of breeding concentrations is related to the grass vegetation. Agriotes tauricus Heyden, 1882. Body size – 9–12 mm. Flat, the pronotum in front of the posterior edges is narrowed. The largest width of the middle segments of the antennae is slightly less than their length. Brown, the abdomen, the antennae and legs are lighter; the upper part is covered in silky hairs (Bey-Bienko, 1965). Distrib- uted in highland Crimea, the Caucasus, Ciscaucasia. The beetles look similar to A. gurgistanus, but differ by gloss and thin silky indumentum. Larvae can be found in Ukraine only on soils of tilled and meadow areas in the mountain regions of Crimea, where they form concentrations of 5 specimens per 1 m2 and cause damage to agricultural crops in a similar way to wireworms of the same genus. Agriotes ustulatus Schaller, 1783. Body size – 8–13 mm. All intevals between the puncture striae on the elytra are similarly hairy, of same color. The second segment of the antennae is notably shorter than the fourth. The punctures on the epipleura of the prothorax are significantly larger compared to the pronotum, V. I. Rusynov, V. O. Martynov, T. M. Kolombar 95 more flat, very densely arranged. Black-brown with silky yellow hairs, the elytra are yellow-brown, the antennae and legs are light. Distributed in Central and South Europe, Algeria, Tunisia, in the south part of the European part of Russia, Ciscaucasia, the Volga region (Furlan, 1998). In Ukraine, the zone of mass distribution and significant damage includes Zakarpatia Oblast, the west and central forest-steppe zone. The larvae are abundant in tilled fields on grey forest soils and leached chernozems. Completely absent on sod-podzolized soils. One of the most harmful pests of seeds, seedlings of agricultural plants and root-tuber crops in forest-steppe zone of Ukraine. Melanotus brunnipes Germar, 1824. Body size – 11–15 mm. The pronotum has no elevated midline, but is smoothened at the front edge. The pronotum has equal length and width, its sides are insignificantly rounded. The terminal sternum of the abdomen is impressed on the sides, so its middle part seems raised, it is cut on the apex and densely covered with hairs. Black, the antennae and legs are brown; in yellow indumentum. Distributed in Asia Minor, in the south of the European part of Russia, the Caucasus (Kabalak and öZbek, 2018). In Ukraine, inhabits forest-steppe and north part of the steppe zone, is especially abundant in the zone of grey forest soils and degraded chernozems in the forest-steppe. The greatest damage is caused by of third and fourth year larvae, younger larvae prefer decomposing root tubers and seeds. Larvae of this species are among the most harmful pests. Common on chernozem soils, in some places, occurs in large amounts in tilled lands in the forest-steppe. Prefer soils of heavy metal compound.

Family Scarabaeidae

Subfamily Cetoniinae Cetonia aurata Linnaeus, 1758. Body size – 15.0–20.5 mm. The dorsal surface is covered in scattered hairs, metallic-glossy, green, golden-green, copper-red, dark-violet-blue, blue, black; the ventral 96 Chapter 2. Coleoptera pests of stored food supplies and field crops surface is copper-red. The elytra bear transversal white spots. Dis- tributed in Europe, West Asia, China, Mongolia, the European part of Russia, except the tundra, Siberia, the Caucasus, Kazakhstan, Uzbekistan, (Englund, 1993). Distributed throughout Ukraine, but sporadically in the south. The beetle harms flowers of fruit and decorative plants, field and garden crops. Tropinota (Epicometis) hirta Poda, 1761. Body size – 8.4–13.6 mm. The elytra have white or yellow-white, sometimes disappearing, spots. Black, mat, the upper part is covered in long white hairs. Distributed in Central and South Europe, North Africa, West Asia, the European part of Russia, the Caucasus, Kazakhstan. Distributed throughout Ukraine, but causes more damage in the steppe zone and in Crimea (Subchev et al., 2011). Beetles gnaw flowers of fruit trees, garden roses, rose (Rose), whitebeam (Sorbus), shadbush (Amelanchier), almond (Prunus dulcis), lemon, mandarin, grape (buds, ovaries, young leaves), buckeye (Aesculum), Viburrum, Syringa, privet (Li- gustum vulgare), Sambuculum, golden currant (Ribes aureum), black currant (Ribes nigrum) (young leaves and flowers), oleaster (Eleaeg- nus), peony (Paeonia), poppy, radish (Raphanus raphanistrum subsp. sativus), Eruca, rape, mustard, cabbage (seeds), beet (transplants), flax, Rheum, cucumber, water melon, melon, pumpkin, kenaf, cotton plant, castor bean, strawberry, sainfoins, pea (leaves, seedlings), clover, vetches, broad bean, soya, wild bean (Phaseolus), lucerne, chick pea (Cicer arietinum), sunflower, Carthalmus, tomato, Bellis, Iris, tulip, ears of rye, wheat, barley, panicle of panic grass, maize and other plants. Oxythyrea funesta Poda, 1761. Body size – 8.9–13.7 mm. The pronotum has no fringe on the sides, has six white spots positioned in two longitudinal rows. The upper part is covered in long hairs, glossy, black-green with bronze shimmer. The elytra bear numerous white spots. The abdomen of females has longitudinal rows of white spots on the sides, which males have also in the middle. Distributed in Southern and partly Central Europe, west regions of the European part of Russia, the Caucasus, Kazakhstan (Tamutis and Dapkus, 2013). Distributed throughout Ukraine. Beetles gnaw V. I. Rusynov, V. O. Martynov, T. M. Kolombar 97 flowers of forest species, fruit trees, decorative plants, field and garden crops.

Subfamily Dynastinae Pentodon idiota Herbst, 1789. Body size – 15.8–25.3 mm. The forehead has one tubercle. The pronotum is much more narrowed in the front than at the base, the widest park is behind the middle. Black, mat. Distributed in Albania, Austria, Bulgaria, Croatia, Cyprus, the Czech Republic, European part of Turkey, France, Hungary, Crete, Macedonia, and the Near East, North Af- rica, Romania, Slovakia, Spain, south part of Russia, the Caucasus (Bunalski et al., 2014). Distributed throughout Ukraine, except the zone of Polesia, damage is most notable in Dnipropetrovsk and Zaporizhia Oblasts. The beetles damage stems at the bases or root collar of many cultivated plants and wild-growing species of plants: maize, sunflower, beet, cabbage, radish (Raphanus sativus var. radicula), rape, buckwheat, Rheum, watermelon, cucumber, melon, pumpkin, kenaf, castor bean, Sesamum, soya, peanut, wild bean, Humelum, canabis, safflower, hyacinth bean (Lablab purpureus), lucerne, sainfoins, poppy, tomato, tobacco, potato, sage, mint, wheat, panic grass, sorghum; can cause damage in plant nurseries, eating seedlings of apple, pear, peach, cherry, sour cherry, cherry plum, mulberry, Persian walnut (Lodos, 1981). Larvae harm roots of maize, sunflower, beet, Carthalmus, wheat, sorghum, onion, potato, grapevine, red and black currant, gooseberry (Ribes uvac- rispa), barberry, apple, pear, apricot, cherry plum, oleaster, fennel, dragonhead (Dracocephalum), lavender and other plants.

Subfamily Melolonthinae Anoxia pilosa Fabricius, 1792. Body size – 17.5–26.5 mm. The pygidium has a groove at the apex. The symmetrical spots on the pronotum are small, the posterior spots are often absent. The sterna of the abdomen have sharply distinct triangular white spots. The hairs on the elytra are small, uniformly scattered, form no patches. The apex of the propygidium bears the same small hairs as the pygidium. Black- 98 Chapter 2. Coleoptera pests of stored food supplies and field crops brown, the elytra are slightly lighter; sometimes partly or completely brown-red. The upper part has dense grey hairs, while the pronotum has long protruding white hairs. Distributed in the European part of Russia, the Caucasus, Kazakhstan. Distributed throughout Ukraine, except Polesia. Larvae damage grain crops (Dekhtiarev, 1929). Melolontha hippocastani Fabricius, 1801. Body size – 20.5–29.0 mm. The pygidium is vertical, among males is narrowed on the apex into a short projection with enlarged apex; pygidium of females is shorter, sometimes undeveloped. Red-brown, vertex, nape, scutellum, epipleura of the elytra, pygidium and the lower part are black. The color of the upper side and legs ranges red-brown to black. Distributed in East Europe, the European part of Russia, Siberia (Ruther et al., 2000). Distributed throughout Ukraine, causing greatest damage in the forest and forest-steppe zones. Beetles gnaw leaves, birch and other trees and shrubs (similarly to M. melolontha). Vora- cious consumer of the needles of larches (Larix) and inflorescence of pine, more rarely – needles of pine, does not consume leaves of ash. Larvae damage roots of the same fruit and tree plants as larvae of M. melolontha, causes great damage, especially in plant nurseries and young plantations (Švestka, 2008). Also damages roots of many field and garden crops: beet, grasses, bean crops, potato, tobacco, carrot, poppy, cabbage, rape, cucumber, melon, water melon, pumpkin, sunflower, hop, buckwheat, Rheum, mallow, flax, Ricinus, sage, mint, onion, garlic, Asparagus and other. Damages roots of grain crops. Melolontha melolontha Linnaeus, 1758. Body size – 22.5–31.5 mm. The pygidium is more gently sloping, with small appressed hairs, which are long and raised only on the sides and the apex, with long (shorter in females) process. The sides of the pronotum bear less dense hairs, which do not cover the main surface. Black, the elytra, pygidium, legs and antennae are red-brown, the elytra and femora are sometimes black (Švestka, 2010). Distrib- uted in Europe, except the north of Scandinavia, Iberian Peninsula, South Italy and Greece, in the west of the European part of Rus- sia (Reinecke et al., 2002). Distributed throughout Ukraine, causes greatest harm in the forest and the forest-steppe zones. V. I. Rusynov, V. O. Martynov, T. M. Kolombar 99

The beetles gnaw leaves of oak, beech (Fagus), different willows, poplars, aspen, maple, buckeye, field elm (Ulmus minor), alder (Al- nus), lime tree (Tilia), birch, Persian wallnut, hazel (Corylus), rose, black locust and Siberian peashrub, gooseberry, grape, staghorn su- mac (Rhus typhina), buckthorns (Frangula), black elderberry and red alderberry (Sambucus racemosa), Tatarian honeysuckle, larch needles ; rarely eat spruce and pine needles, also there were recorded cases of significant damage caused by the beetles to leaves, flowers and ovaries of fruit trees, especially sour cherry, cherry and apple trees. Causes no damage to leaves of ash tree and Syringa. Larvae harm grain crops, gnaw roots of birch, oak, hazel, maple, ash, black locust (Robinia pseudoacaci), buckeye, Persian walnut, all fruit trees, berry shrubs, rose, Swida, lilac, privet, Tatarian honeysuckle, sil- verberry, Viburrum, barberry, grapevine, herbaceous plants, grain crops, bean crops, potato, tobacco, poppy, cabbage, beet, rape, cucumber, melon, water melon, pumpkin, carrot, sunflower, buckwheat, Rheum, mallow, flax, Ricinus, sage, mint, onion, garlic and Asparagus. The greatest damage is caused to sugar beet, potato, strawberry, and also forest and fruit trees, especially in plant nurseries and young plantations. Amphimallon assimile Herbst, 1790. Body size – 11.2–14.0 mm. The spurs on the internal edge of the protibiae are positioned opposite the middle tooth of their outer edge which bears three teeth. The disk of the pronotum bears thin protruding and sometimes also appressed hairs. The pronotum has small, very dense, punctures. The elytra have short appressed hairs. The scutellum is significantly transversal. The pronotum is almost mat, with small and dense punctures, in protruding and appressed hairs. Light colored, red- brown or dark-brown. Distributed in the south-west part of Europe- an Russia (Gavrilovic and Curcic, 2010). In Ukraine can be found in western Oblasts, and then further to the east up to Vinnitsa, Odessa and Mykolaiv Oblasts in the east. Larvae gnaw roots of cultivated plants, similar to larvae of A.solstitialis. Amphimallon altaicus Mannerheim, 1825. Body size – 12.8–15.5 mm. The pronotum is covered in scattered punctures of 100 Chapter 2. Coleoptera pests of stored food supplies and field crops average size, the males have long, protruding, and the females – shorter appressed hairs. The scutellum is semi-rounded. The elytra of males have rather short hairs, females are almost bare. Black or black-brown, the elytra sometimes are red-brown. Distributed in the European part of Russia, the Caucasus, Siberia, Kazakhstan, on the Balkan Peninsula. In Ukraine, can be found in Lviv and Donetsk Oblasts, but very sporadically. The larvae harm the roots of field crops (Dudarova and Abdurahmanov, 2009). Amphimallon solstitiale Linnaeus, 1758. Body size – 13.8–19.0 mm. The spur on the internal edge of the protibiae is positioned opposite the groove between the upper and the middle teeth of the outer edge (if three teeth are present) or opposite the groove between the teeth (if two teeth are present). The disk of the pronotum, besides fine hairs, also has large setae. The external edge of the protibiae of males has 1–2, rarely 3 teeth, females – 3 teeth. The elytra have large ribs. Pale-yellow, the abdomen, spot in the middle of the pronotum, divided by a longitudinal stripe, often dark. The pronotum of males has long appressed and protruding, and females – short appressed hairs (Montreuil, 2000). Distributed in Europe, Asia Minor, North Iran, Mongolia, China, the European part of Russia, Siberia, the Caucasus, Central Asia. Distributed throughout Ukraine. In forest and forest-steppe zones, beetles gnaw leaves of fruit trees, raspberry, pine needles Larvae damage roots of pine (two years old seedlings), maples, ash trees, fruit trees, prickly wild rose (Rosa acicularis), black locust (Robinia pseudoacacia) and Siberian peashrub (Caragana arborescens), locust (Gleditsia), European spindle (Euonymus europaeus) and Euonymus verrucosus, black elder (Sambucus nigra), barberry, Viburrum, currant, gooseberry, hop (Humulus), Persian walnut, grape, stalk of poplar, roots of poppy, cabbage, rape, buckwheat, Rheum, flax, beet, cucumber, melon, water melon, pumpkin, cotton plant, mallow (Malva), castor bean, pea, wild bean, soya, peanut, cannabis, carrot, valeriana, sunflower, chry- sanths (Chrysanthemum), safflower, tobacco, tomato, sage, mint, lavender, Lallemantia, potato, onion, garlic, Asparagus, chufa, wheat and other grain crops, maize and Sudan grass. V. I. Rusynov, V. O. Martynov, T. M. Kolombar 101

Holochelus (Miltotrogus) aequinoctialis Herbst, 1790. Body size – 13.5–18.5 mm. The capitulum of the males’ antennae is significantly bent and longer than the pedicel. The males’ pronotum has dense small punctures and long protruding hairs, females’ pronotum has dense punctures and shorter hairs. Glossy, brown-red. Distributed in South- East Europe, Asia Minor, the European part of Russia, Kazakhstan, the Caucasus (Gavrilovic and Curcic, 2010). Distributed throughout Ukraine, most harmful in the steppe zone. Larvae gnaw roots of cabbage, rape, swede, turnip rape (Brassica rapa), buckwheat, Rheum, flax, mallow, beet, soya, peanut and other bean crops, Ricinus, hop, carrot, sunflower, safflower, potato, tobacco, eggplant (Solanum mel- ongena), tomato, maize, wheat and other grain crops, onion, grape, currant, gooseberry, rose, seedlings and saplings of fruit trees, oak, maple, ash trees, black locust and Siberian peashrub, Tatarian honeysuckle (Lonicera tatarica), European spindle and others. Holochelus (Miltotrogus) tauricus Blanchard, 1850. Body size – 13–16 mm. The capitulum of the males’ antennae is slightly bent, of the same length as the pedicel; the sixth and seventh segments of the antennae are transversal. The pronotum has small similar punctures and long (shorter among females) protruding hairs. Glossy, red-yellow and brown-red. Endemic species to the Crimea (Martynov, 2010). Larvae cause damage in a similar way to larvae of M.aequinoctialis. Holochelus (Miltotrogus) vernus Germar, 1823. Body size – 16–20 mm. The pygidium has similar punctures. The appressed hairs on the pronotum are only slightly shorter than the protruding; hairs of females are much shorter compared to males. The pronotum is mat, in very small and dense punctures. Insignificantly glossy, dark brown-red. Distributed in the south of the European part of Russia. Distributed throughout Ukraine except the Crimea (Vovk et al., 2016). Larvae cause damage in a similar way to the larvae of Miltotrogus aequinoctialis. Monotropus nordmanni Blanchard, 1851. Body size – 10.7–12.5 mm. The head, pronotum, elytra, at least at the base, and pygidium is covered with long protruding hairs. Yellow, the forehead, the disks of the pronotum and the scutellum are brown, the sides of 102 Chapter 2. Coleoptera pests of stored food supplies and field crops the elytra have narrow brown margin. Distributed in the Mediter- ranean, the south of the European part of Russia (Coca-Abia, 2003). Distributed throughout Ukraine, except the Crimea. Lives in sands, larvae gnaw roots of plants, harms grapevines, seedlings and saplings of tree species. Polyphylla fullo Linnaeus, 1758. Body size – 26–36 mm. The protibiae of males have two teeth, and of females – three. White scales form longitudinal stripes on the pronotum, the elytra bear white marble pattern of fusing spots. The base of the pronotum has no fringe. The elytra have distinct white or yellow-white scale spots, between which only rare single scales are scattered. Brown- black or reddish-brown. Distributed in Central and South Europe, south and middle belt of the European part of Russia. Distributed throughout Ukraine on sandy soil, massively abundant and causes the greatest damage in forest-steppe and steppe zones, especially in lower Dnipro sands (Dekhtiarev, 1929). The beetles gnaw needles of pine, leaves of beech, poplar, black locust, etc. Larvae cause significant damage by gnawing roots of young tree and shrub species – pine, oak, grape (especially stalks), fruit trees, berry shrubs; harms roots of cabbage, rape, poppy, beet, buckwheat, Rheum, bean crops, Ricinus, sunflower, carrot, potato, tobacco, sage, mint, onion, garlic, Asparagus, grain crops, maize. Especially harmful for young pine plantations, plant nurseries and vineyards on sands. Rhizotrogus aestivus Olivier, 1789. Body size – 13.0–17.2 mm. Yellow, middle of the pronotum has a brown longitudinal stripe, the elytra along the elytral suture and the side margins are brown (Bey- Bienko, 1965). Distributed in Central and South Europe, West Asia, the south of the European part of Russia. In Ukraine is distributed to the north to Khmelnitsky, Vinnitsia, Kyiv, and Sumy Oblasts. Larvae gnaw roots of field and garden crops and tree plants. Can cause significant damage in plant nurseries. Maladera holosericea Scopoli, 1772. Body size – 7.0–8.2 mm. Black-brown, sometimes reddish, mat, with dove-colored tinge. The antennae and legs are brown-reddish. Distributed in Central and Eastern Europe, the European part of Russia, the Caucasus, Siberia, V. I. Rusynov, V. O. Martynov, T. M. Kolombar 103

Kazakhstan, Central Asia (Ahrens, 2006). Can be found throughout Ukraine on sand. Way of life similar to S. brunnea. Beetles gnaw leaves and buds of field and garden crops, larvae - roots of herbaceous plants, tree seedlings and saplings. Serica brunnea Linnaeus, 1758. Body size – 8–10 mm. Oblong, mat, brown-reddish, with slight whitish silky shimmer. Distributed in Europe, the European part of Russia, Siberia, Kazakhstan. Dis- tributed throughout Ukraine, except the Crimea, but sporadically in the south. Beetles gnaw leaves of trees and shrubs, larvae – young roots of field crops and seedlings of tree plants (Eyre et al., 2002).

Subfamily Rutelinae Anisoplia (Anisoplia) agricola Poda, 1761. Body size – 10.5–13.0 mm. Skinlike border of the elytra reaches the posterior edge of the III sternum of the abdomen. The pronotum is covered in short protruding hairs. Black, with metal-green shimmer, the elytra are reddish, yellow or brown, of same color or with black suture, spot at the scutellum and sash at the middle, or entirely black (Mico et al., 2001). Distributed in middle and south belt of the European part of Russia, the Caucasus, Kazakhstan. Distrib- uted throughout Ukraine, more often in Polesia and northern part of the forest-steppe, and also in the pre-mountain and mountain area of the Crimea. Beetles damage immature seeds of wheat, rye, barley, foxtail millet, larvae – tubers of potato, roots of beet, grain crops. Anisoplia (Anisoplia) brenskei Reitter, 1889. Body size – 9.3–11.0 mm. Large claw of the front legs of males are thicker, less narrowed to the apex which is notably cut short. The pronotum is parallel, narrows only in the front, has an insignificant grooves on the sides in front of the posterior angles. The upper part is yellow- grey, the lower part bears white hairs. Distributed in the middle and south of the European part of Russia (Pavlov and Ruchin, 2013). Distributed throughout Ukraine in the Left-Bank forest-steppe, among forbs of the steppe, in the premountain and mountain areas of Crimea. On rye ears usually eats anthers, more rarely – ovaries. 104 Chapter 2. Coleoptera pests of stored food supplies and field crops

Anisoplia lata Erichson, 1847. Body size – 12–15 mm. Skinlike border of the elytra hardly reaches the basis of the IV sternum. The pronotum is bare. The elytra are of one color – brown-red, brown or black. The pronotum is narrowed from the basis to the front, its sides are almost straight in front of the posterior angles. The patch of hairs near the scutellum on the elytra of males is not obvious, and the lateral swollen part of females reaches the last third of the elytra. The abdomen bears scattered hairs. Rather glossy, black, the head and the pronotum have greenish shimmer. The elytra are red- brown, brown or black-brown. Distributed in South and South- East Europe, European part of Russia, Moldova (Mico et al., 2001). In Ukraine, is distributed in the north up to Vinnytsia, in the east up to Cherkassy Oblast. Causes the most significant damage in the steppe zone of the Right Bank. The beetle damages immature seeds of wheat, rye, and barley. Anisoplia segetum Herbst, 1783. Body size – 8.0–12.5 mm. The elytra bear scattered hairs, long and protruding at the base, the females have an insignificant lateral swollen part. The head and the pronotum have scattered protruding hairs. Black, with bronze or green gloss, the elytra are brown-yellow, females have black spot at the scutellum. Distributed in the south of the European part of Russia, the Caucasus, Kazakhstan (Blintsov, 1986). Distributed throughout Ukraine, significant damage observed in the steppe zone and the Crimea. Beetles gnaw anthers, and also ovaries and seeds at the beginning of ripening on the ears of rye, wheat, and barley. The larva eats roots of beet, sunflower, chufa, tobacco, seedlings of apple tree, pear, plum, tubers of potato. Anisoplia tempestiva Erichson, 1847. Body size – 10.8–13.5 mm. The lateral edge of the elytra has short thick setae. The dorsal surface is glossy, the elytra are bare or with a hairy spot near the scutellum, the females have a well developed lateral swollen part; if the elytra are light colored, of one color, then no apical infuscation is present. The legs are of moderate length. The skinny fringe of the elytra is long, stretches from the elytral suture angle to the place of widening of the lateral edge. The sides of the pronotum of males are parallel, V. I. Rusynov, V. O. Martynov, T. M. Kolombar 105 narrowed only in the front. Metal-green, the elytra are brown, rarely with black pattern (Bey-Bienko, 1965). Distributed in Palestine. In Ukraine can be found in Zakarpattia Oblast. The beetles gnaw immature grains of crops. Anisoplia zwicki Fischer von Waldheim, 1824. Body size – 10.2 – 13.0 mm. The pronotum of males and females narrows from the base to the front, with straight, slightly grooved lateral edges in front of them. The elytra of males and females have an obviously hairy area near the scutellum, the lateral swollen part of females is only slightly behind the middle of the elytra. The abdomen is covered with dense grey hairs. Almost mat, black, sometimes with green shimmer, the elytra are brown-red or black (Baraud, 1991). Distrib- uted in the south and south-east of the European part of Russia, Ciscacausia, Kazakhstan. In Ukraine, distributed in Luhansk and Donetsk Oblasts and in Crimea. More often on wild grasses, especially couch grass; sometimes harms grain of wheat, rye, and barley, similarly to A. austriaca. Anisoplia (Autanisoplia) austriaca Herbst, 1783. Body size – 13–16 mm. The abdomen and femora have short appressed hairs. The lateral edge of the elytra bears short thin setae. Bare on the dorsal surface, black, with metal-green shimmer, the elytra are red-brown, females always have black spot near scutellum, whereas males have this only in the south of their range. Distributed in West Asia, the south of the European part of Russia, the Caucasus (Ozdar, 2002). Distributed throughout Ukraine, except north-west regions of Polesie. The most significant damage is caused in south of the forest-steppe and steppe, south from the line between Vinnit- sia, Kyiv, Poltava and Kharkiv Oblasts. The beetle gnaws out seeds of grasses during the period of milky ripeness, and pushes solid grasses to the ground; particularly damaging to wheat, rye and barley, feeds on grains of wild grasses - different species of couch grass and other species. Larvae gnaws roots of rye, wheat, maize, chufa, beet, sunflower, potato, tobacco, seedlings of apple tree, pear, plum, cherry, mulberry and other. 106 Chapter 2. Coleoptera pests of stored food supplies and field crops

Anomala errans Fabricius, 1775. Body size – 9.5–13.4 mm. The pygidium bears very short hairs, which are longer on the on sides and the apex. The pronotum has small similar punctures, with a fringe on the posterior edge, interrupted in the middle. Yellow with black; sometimes black color dominates or dominates over yellow. Distributed in the south of the European part of Russia, Kazakhstan, Uzbekistan (Saypulaeva, 2015). In Ukraine, is found in the south from Kyiv, Poltava, Kharkiv up to Crimea, on sands. Beetle does not eat, the larvae gnaw roots of field, garden and forest crops. Phyllopertha horticola Linnaeus, 1758. Body size – 8.4–11.0 mm. Metal-green, blue or black, the elytra are brown, sometimes black. Distributed in the European part of Russia, the Caucasus, Siberia, Kazakhstan, Kyrgyzstan. In Ukraine, can be found in Polesie, the forest-steppe, and in the northern regions of the steppe zone. The beetle gnaws flowers and young leaves of peony, barberry, cabbage, rape, cucumber and other cucurbits (Cucurbitaceae), beet, currant, gooseberry, apple, pear, plum, cherry, raspberry, garden rose, rose, blackberry, lucerne, clover, lupin, pea, vetches, broad bean, hop, cannabis, mulberry, sea buckthorn (Hippophae), vinegrape, Viburrum, Persian walnut, lilac, sunflower, Scorzonera tau-saghyz, iris, maize and other plants (Raw, 1951). Larva gnaws roots of grain grasses, beet, clover, tree seedlings.

Family Geotrupidae

There are 250 species in Ukraine, of which 70 are pests of agriculture and forestry, and 8 are especially harmful: Anisoplia austriaca, Melolontha melolontha and Melolontha hippocastani, Polyphylla fullo, Anoxia pilosa, summer chafer (Amphimallon solstitiale), Mil- totrogus aequinoctialis, Lethrus apterus. Development takes place in the soil, burrows of mammals (in the bedding of nests or food stores), accumulation of plant remains, rotten wood, tree hollows, stumps, anthills, manure, rarely – in corpses of animals. Some species, for example, earth-boring dung beetles, prepare food for their progeny in form of balls, pears, V. I. Rusynov, V. O. Martynov, T. M. Kolombar 107 sausages, which are placed in the soil; Lethrus apterus drags gnawed parts of plants into its burrow, into special cells. The fertility of females is low (40–50 eggs for summer chafers), and especially low in species which take care of their progeny (Lethrus apterus lays 8–10 eggs, maximum 20 eggs). Adult beetles feed on plant food, plant remains, manure, rarely corpses of animals, some do not eat at all (Anoxia pilosa, Rhizotrogus vermis, Miltotrogus aequinoctialis). Natural enemies of chafers are some parasites – nematodes (Mer- mitidae), Gamasina mites, tachina flies (Dexia and other), scoliid wasps (its larva is an external parasite of the chafer), and also predators – ground beetles, robber flies (Asilidae), spiders, toads (particularly Pelobates fuscus), lizards, very many birds, insectivorous mammals. Some species cause harm during the larval and imago (chafers) stages, others only at the larval stage (A. pilosa, Rhizotrogus and Miltotrogus), and others – only at the imago stage (Oxythyrea funesta) flower chafers (Cetoniinae). The beetles gnaw leaves and immature fruits (summer chafers), flowers (O. funesta, flower chafers), immature seeds of grasses (An- isoplia), gnaw out tunnels in stems (flower chafers) or at their bases (Pentodon idiota), damage mature fruits (flower chafers). Lethrus apterus eats buds, leaves, shoots of plants, out of which it prepares food for its progeny. Larvae of species which live in soil gnaw roots, tubers of herbaceous and tree plants, often causing their death. Lar- vae of older ages of large species (summer chafer, Polyphylla fullo, A. pilosa) are especially harmful, because they gnaw through large roots and need large amounts of food. Some species are carriers and intermediary hosts of helminths – parasites of domestic animals (earth-boring dung beetles, larvae of flower chafers and summer chafers).

Subfamily Lethrinae Lethrus apterus Laxmann, 1770. Body size – 15–24 mm. Black, slightly glossy, sometimes slightly bronze, with a bluish shimmer. The lower processes of the mandibles of males are identical, 108 Chapter 2. Coleoptera pests of stored food supplies and field crops small, identically bent inside, with diagonal longitudinal furrow (Bey-Bienko, 1965). Distributed in Central Europe, the Eastern Medi- terranean, the eastern part of European Russia. Distributed throughout Ukraine, except Zakarpatia Oblast and Crimea (Bunalski et al., 2014). Settles in fields, steppe slopes, railway embankments, gullies, dry ravines, on the sides of roads, in undisturbed areas with consoli- dated soil (Rosa et al., 2017). Absent in sands and swamps. Beetles winter at the depth of up to 50 cm. From mid April to mid June, the beetles make temporary burrows on slopes, which extend at the angle of 25–30 ° down to 15–18 cm, where they live, hide in the night and in cases of threats. Active on warm sunny days. After mating, the male and female organize a mutual burrow with offshoots and together prepare food for the progeny to come. The burrow comprises a slant- ing upper bend of 18–27 cm depth, which leads to a vertical tunnel to depth of 54–63 cm, sometimes down to 72 cm; along this bend, the female makes a 3 cm long and 2 cm wide chamber, in the wall of which, the egg is laid. After the egg is laid, the chamber is filled with fragmented green parts of plants, and then patched up with earth. On average, the female lays 8–11 eggs, maximum 20. The egg stage lasts 10–12 days. After the larva emerges from the egg, it feeds on food which the parents have prepared in the chamber, lives three weeks, moults three times and after the terminal moult, transforms into a pupa, out of which after 12–14 days the beetle hatches, which stays in the chamber over the winter, and exits it only in spring. Larvae of Lethrus apterus can be infested with Oospora destructor (Metschnikoff) Delacroix 1893, bacteriosis and destroyed by larvae of robber flies (Asilidae), Hister beetles (Margarinotus bipus- tulatus Schrank, 1781) and skin beetles (D. lardarius). The beetles are consumed by rooks, European rollers, lesser grey shrikes. The beetles are parasitized by Acari.

Family Silphidae

There are 900 species of large carrion beetles around the world, 30 in Ukraine. V. I. Rusynov, V. O. Martynov, T. M. Kolombar 109

The beetles are various in shape, color and sizes (up to 6 to 40 mm long). The antennae are usually clavate, the front coxae are protruding. The elytra often do not cover the upper part of the abdomen. The legs have five segments. The most typical form of the body of larvae is woodlouse-like. Most species feed on corpses of animals, some are predators or her- bivores. The latter eat the trama of edible mushrooms, sprouting seeds, young shoots and leaves of grain, technical, garden and forage crops. Subfamily Silphinae Aclypea opaca Linnaeus, 1758. Body size – 9–12 mm. The upper part is covered in dense appressed grey-yellow hairs, the inter- sections of the elytra have no coarse wrinkles. The species is distinguished from its relatives by its thickened head without a joint between the head and the thorax and deeply grooved upper lip. Dis- tributed in North America, Austria, Belarus, Belgium, Great Britain, Bulgaria, the Czech Republic, Denmark, Estonia, Finland, France, Germany, Hungary, Ireland, Italy, Latvia, Liechtenstein, Lithuania, Luxemburg, Macedonia, Moldova, the Near East, Norway, Poland, Slovakia, Slovenia, Spain, Sweden, Switzerland, Netherlands, the European part of Russia, Siberia (Aleksandrowicz and Komosiński, 2005). In Ukraine can be found in Polesia and the forest-steppe (Brygadyrenko, 2015). Damages maize and other grain crops, beet, rape, lucerne. The beetles sometimes eat the trama of edible mushrooms (penny bun, Russula) and fruits of strawberry. Aclypea undata Muller, 1776. Body size – 11–15 mm. The dorsal surface is almost bare, the intervals between the longitudinal keels of the elytra are coarsely and irregular wrinkled-punctured. Black, almost mat. Distributed in Albania, Austria, Belarus, Belgium, Great Britain, Bulgaria, Croatia, the Czech Republic, Denmark, Estonia, Finland, France, Germany, Greece, Hungary, Italy, Latvia, Lithuania, Luxemburg, Macedonia, Moldova, the Near East, Poland, Portugal, Romania, Slovakia, Slovenia, Spain, Sweden, Switzerland, the Neth- erlands, the European part of Russia, Central Asia (Dekeirsschieter 110 Chapter 2. Coleoptera pests of stored food supplies and field crops et al., 2011). Beetles and larvae harm grain (barley, maize, panic grass), technical (beet, sunflower, potato) and forage (clover, lucerne) crops, flesh of strawberry and tomato, fallen fruits in gardens. Phosphuga atrata Linnaeus, 1758. Body size – 10–16 mm. The head is significantly stretched. The elytra have sharply distinct ribs. Black or red-brown, glossy. The elytra between the ribs bear dense punctures. Distributed in Albania, Austria, Belarus, Belgium, Great Britain, Bulgaria, Croatia, the Czech Republic, Denmark, Estonia, Finland, France, Germany, Hungary, Ireland, Italy, Latvia, Leichten- stein, Lithuania, Luxembourg, Macedonia, Moldova, the Near East, Norway, Poland, Portugal, Romania, Slovakia, Slovenia, Spain, Swe- den, Switzerland, the European part of Russia, Siberia, Japan (Suzzo- ni, 1972). Damage to beet has been observed. Has been mentioned as a pest of grain crops, rape, however these data should be checked. Silpha carinata Herbst, 1783. Body size – 12–23 mm. The eighth segment of the antennae is significantly longer than the ninth. The lateral edge of the elytra is very wide, especially in the front, significantly explanate. The elytra have high glossy ribs, with no preapical tubercle (Madge, 1980). Black-brown, more rarely black or brown-red. Distributed in Albania, Austria, Belarus, Belgium, Great Britain, Bulgaria, the Czech Republic, Denmark, Estonia, Finland, France, Germany, Hungary, Italy, Latvia, Lithuania, Luxembourg, Moldova, the Near East, Norway, Poland, Romania, Slovakia, Slove- nia, Sweden, Switzerland, the Netherlands. Silpha obscura Linnaeus, 1758. Body size – 13–17 mm. The middle coxae are close to one another. The head has a joint between the head and the thorax. The elytra bear longitudinal keels. The punctures on the elytra are simple, with no glossy grains. Has a row of smaller punctures along the ribs of the elytra on each side; the punctures on the sides of the elytra are two times smaller than in the internal intervals. The ribs are weak, but sometimes all (or only internal) are significantly raised and more glossy. The sides of the elytra are uniformly rounded (Heymons and Lengerken, 1926). Black, mat. Distributed in Albania, Austria, Belarus, Belgium, Great Brit- ain, Bulgaria, Croatia, the Czech Republic, Denmark, Estonia, Fin- V. I. Rusynov, V. O. Martynov, T. M. Kolombar 111 land, France, Germany, Greece, Hungary, Italy, Latvia, Luxemburg, Macedonia, the Near East, Norway, Poland, Romania, Sardinia, Sic- ily, Slovakia, Slovenia, Spain, Sweden, Switzerland, the Netherlands, the European part of Russia, Siberia, Central Asia. Is distributed throughout Ukraine. The larvae harm beet (leaves, rarer wheat, rye, barley, maize, sunflower, turnip rape, rape and cabbage plants. The beetles sometimes eat fruits of strawberry and tomatoes.

Family Staphylinidae

The fauna of Staphylinidae or rove beetles comprise 25.000 species around the world, including 900 in Ukraine. The body of the beetles is narrow, the elytra are usually short, covering only the first two abdominal terga. Usually has wings which are folded in longitudal and trasversal orientations and are almost entirely covered by the elytra. The legs are of running type, the antennae are filiform or clavate. The abdomen comprises 6–7 distinct segments, very flexible, and when beetles run, they bend it upward. The larvae have long oligopod body. The beetles and larvae are mostly predatory, live in various decomposing organic matter, under rocks, tree bark, mosses, mushrooms, in coastal sand, nests of different animals, and sometimes flowers.

Subfamily Oxytelinae

Coprophilus (Coprophilus) striatulus Fabricius, 1793. Body size –5.5–6.5 mm. The pronotum has saw-toothed edges, the disk bears longitudinal impressions and middle furrow. Brown-black, glossy. Distributed in Austria, Belgium, Great Britain, Croatia, the Czech Republic, Denmark, Estonia, Finland, France, Germany, Hungary, Italy, Latvia, Lithuania, Moldova, Norway, Poland, Roma- nia, Slovakia, Sweden, the Netherlands, the European part of Russia (Hoebeke, 1995). In Ukraine, can be found in Polesia. Under decomposing remains, in manure, some species consume fleas in the 112 Chapter 2. Coleoptera pests of stored food supplies and field crops burrows and shelters of mammals. Beetles eat trama of different mushrooms. Mentioned as a pest of sprouting seeds of maize and other cultivated plants.

Subfamily Staphylinina Staphylinus caesareus Cederhjelm, 1798. Body size – 17–25 mm. The epimera of the prothorax are absent. Some terga of the abdomen on the sides have large spots of golden-yellowish hairs. Black, the elytra, antennae and legs are brown-red. The scutellum has black hairs. The base of the prontotum has a fringe of yellow hairs. The temples bear yellow hairs (Balog et al., 2008). Distributed in Aus- tria, Belgium, Great Britain, Bulgaria, Croatia, Cyprus, the Czech Republic, Denmark, Estonia, Finland, France, Germany, Hungary, Italy, Latvia, Lithuania, Luxembourg, the Near East, North Africa, Norway, Poland, Portugal, Romania, Slovakia, Slovenia, Spain, Swe- den, Switzerland, the Netherlands. Lives under decomposing vegetative substances, in manure and under rocks. V. I. Rusynov, V. O. Martynov, T. M. Kolombar 113

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Chapter 3. Biological control of beetle pests of stored grain and field crops

V. O. Martynov, V. V. Brygadyrenko Oles Honchar Dnipro National University

Introduction 1

In natural conditions, the main regulating factor of the populations of pests is their natural enemies. Also, natural control includes other biological factors – competition, accessibility of food, and also abiotic factors, among which temperature and moisture are critical. In the struggle against pests, biological control is orientated towards decreasing the pest population through the impact of predators, parasites and pathogens. Biological control can be considered as an element of natural control which involves manipulation of the population of natural enemies by man and can be used for any type of organism, regardless of whether it is a pest or not. There are different definitions of biological control. According to Alston (2011), it is “any activity of a species which reduces the non-favourable impact of another species”. Also, biocontrol can be defined as “study and use of predators, parasites and pathogens for regulating the density of pests” (De Bach, 1964). By contrast to biological control, natural control is not related to human manipulations. The history of biological control dates far back to antiquity. Around two thousand years B.C., the Ancient Egyptians used biological control for the first time, when they began domesticating cats, for they feed on rodents which cause damage to the harvest. The first written records of biocontrol belong to the III century

1 Martynov V. O., & Brygadyrenko V. V. (2019). Biological control of beetle pests of stored grain and field crops. In: Current problems of agrarian industry in Ukraine. Accent Graphics Communications & Publishing, Vancouver, Canada. – P. 134–190. Doi:10.15421/511904 V. O. Martynov, V. V. Brygadyrenko 135

AD when in China, in the territory where the city Guangzhou is located today, a nest of weaver ants (Oecophylla smaragdina (Fabricius, 1775)) was sold for use against such pests as Tessarato- ma papillosa (Drury, 1770). By 1200, the significance of ladybirds (Coccinellidae (Latreille, 1807)) has been recognized as an agent of control against Aphids and Coccoidea. The first description of parasitism of insects was illustrated by van Leeuwenhoek in 1701. Cordyceps fungi which affect owlet moths (Noctuidae) were considered to be the first pathogens of insects by Réaumur in 1726. In 1762, the British and the French successfully introduced the common myna (Acridotheres tristis (Linnaeus, 1766)) from India to Mauritius to use against the red locust (Nomadacris septemfas- ciata (Audinet-Serville, 1883)). In 1776, in Europe, a successful control of common bed bug (Cimex lectularius (Linnaeus, 1758)) was effected using Picromerus bidens (Linnaeus, 1758), a predatory shieldbug. In 1837, Koller for the first time described the conception of “natural control”. In 1850–1887, the conception of biological control developed in the United States. In 1870, Charles Riley for the first time conducted a successful transportation of parasitoids from Kirkwood (Missouri) to other states for biological control of plum curculio (Conotrach- elus nenuphar (Herbst, 1797)). In 1883, the US Department of Agri- culture (USDA) imported Cotesia glomerata (Linnaeus, 1758) from England for the control of the small white (Pieris rapae (Linnaeus, 1758)) common in several states. It was the first intercontinental transportation of parasites. In Europe, projects developed for biological control were first implemented only in the 1800s. Intensification of horticulture during that period led to the search of new methods against insect pests in America as well. The first project on cottony cushion scale (Ic- erya purchasi (Maskell, 1878)), aimed at control of its number, was started in California in 1888, using the vedalia beetle Rodolia car- dinalis (Mulsant, 1850). Since that time, a number of projects have been started, including the project on gypsy moth (Lymantria dispar (Linnaeus, 1758)) in New England (1905–1911). Overall, more than 136 Chapter 3. Biological control of beetle pests of stored grain and field crops

6.000 introductions of over 2.000 insect species have been made for use as control agents against insect pests (Cock et al., 2016). The peak of activity in biological control with 57 different agents created in different places was observed from 1930 to 1940. During the World War II, a steep decrease in activity in biological control occurred, and after the war, it did not become popular due to production of relatively cheap synthetic pesticides. Biological control was revived as the main component of the conception of integrated pest management (IPM) only in the late 1960 s (Kwenti, 2017).

Biological control as a method against pests

Peculiarities of biological pest control The impact of biological control on the diversity of local species can be both positive and negative. The main problem of biocontrol is the impact of the agents on non-target species. Therefore, during the introduction of the control agent, the main requirement is specificity to the host. A potential agent of biological control should undergo a variety of tests and quarantine before being released into the environment, for if an agent affects a broad range of species, it can be highly invasive, causing changes in biodiversity on a large scale. The hardy and prolificGambusia holbrooki (Girard, 1859) which lives in the South-East of the USA has been distributed all around the world for elimination of larvae of mosquitoes and the struggle against malaria. Unfortunately, it has caused great damage to endemic species of fishes and amphibians, as it competes with them for food resources and eats their roe and fry. The short spawning period, high fertility and resistance to salinity have contributed to the wide distribution of Gambusia (Vila-Gispert et al., 2005; Alcaraz & Garcia-Berthou, 2007). Biological control of invasive pests can be conducted by natural inhabitants of the territory of the pests’ distribution. Over time, they adapt to the presence of immigrants and have a negative effect on them through predation or competition, leading to a significant decrease in the number of pests. The birch leafminer (Profenusa V. O. Martynov, V. V. Brygadyrenko 137 thomsoni (Konow, 1886)), brought to North America from Europe, became a significant pest of birch in the 1970s. However, in the 1990s (MacQuarrie, 2008; Soper et al., 2015), its population steeply decreased under the impact of an American parasite Lathrolestes thomsoni (Reshchikov, 2010). Unfortunately, in most cases, local species are not able to control the population of pests sufficiently. For example, in Europe, the invasive horse-chestnut leaf miner (Cameraria ohridella (Deschka & Dimic, 1986)) is attacked by dozens of parasitoids, which causes no significant impact on the level of damage (Grabenweger et al., 2010). The dynamic of populations of invasive insects demonstrates increases and decreases, where their abundance and the damage they reach their peak soon after the invasion, following which their population begins to collapse. One of the reasons for collapse is the activity of local enemies. Kenis & Branco (2017) presume that local natural enemies will affect the invasive insects if they belong to a group of insects which is usually attacked by polyphages or is ecologically and taxonomically closely related to the invader. Over time, local natural enemies start attacking invasive insects and can partly or completely control the pest, but the results are quite unpredict- able and in most cases are insufficient for providing a satisfactory level of control. In some cases, soon after the invasion of a pest, their exotic natural enemies occur, which perform a significant control. For example, parasites of invasive insect pests of Eucalyptus in Mediter- ranean regions, such as Avetianella longoi (Siscaro, 1992) – parasite of longhorn beetle Phoracantha semipunctata (Fabricius, 1775), Closterocerus chamaeleon (Girault, 1922) – parasite of Chalcid wasp (Ophelimus maskelli (Ashmead, 1900)) and Psyllaephagus bliteus (Riek, 1962) – parasite of plant lice (Glycaspis brimblecombei (Moore, 1964)) (Bush et al., 2016). Biological control is one of a few methods against pests which has a constant pattern and which requires no measures after the releasing and distribution of the agents. From a long-term perspective, biological control has a number of advantages. However, the 138 Chapter 3. Biological control of beetle pests of stored grain and field crops assessment of benefits is complicated due to urgency of implementation, complexity of the methods, absence of financial support and low priority given to post-release monitoring. Despite the dif- ficulties, Cock (2015) provide data which shows that the projects of biological control of cassava mealybug (Phenacoccus manihoti (Matile-Ferrero, 1977)) and mango mealybug (Drosicha mangiferae (Stebbins, 1903)) in Africa had the coefficients of costs and benefits of 1: 738 and 1: 808 respectively. Similar results were presented in the studies by Hill & Greathead (2000), Gutierrez et al. (1999), Dahlsten et al. (1998) and others, which demonstrates the obvious advantage of biological control from the perspective of economics. However, it should be mentioned that the development of technology of the control and the subsequent monitoring of the studied ecosystem requires a lot of time.

Risks of biological control In 1980s, concerns arose regarding the ecological safety of agents of biological control due to various side effects on the local biodiversity, which are still voiced today. This problematic was analyzed in the studies by Hajek et al. (2016) and Van Driesche & Hoddle (2017). The main risk related to the introduction of a foreign predator, parasite or pathogen, is its potential effect on local non-target organisms, which leads to the changes in the populations and communities. One of the most known examples is the tachinid fly species (Compsilura concinnata (Meigen, 1824)) introduced to North America for the control of gypsy moth, causing decrease in the native species even today (Elkinton & Boettner, 2012). Side effects can occur as a result of competition with close relative species of predators, for example, competition of ladybugs Harmonia axyridis (Pallas, 1773) and Coccinella septempunctata (Linnaeus, 1758) with native species (Evans, 2004; Roy et al., 2016). The introduction of agents of biological control can have a negative effect as a result of hybridization with related species. Yara et al. (2007) present data showing that Torymus sinensis (Kamijo, 1982) a parasite introduced to Japan as a control agent against the gall wasp (Dryo- V. O. Martynov, V. V. Brygadyrenko 139 cosmus kuriphilus (Jasumatsu, 1951)), caused reduction in the population of T. beneficus (Yasumatsu & Kamijo, 1979) as a result of hybridization. However, competition between these species is also possible. Despite this fact, currently, a relatively low number of introductions have directly affected particular non-target species at the level of population. Indirect impacts are possible, but they are mostly theoretical in character. The last 20 years have wit- nessed the development of international principles, rules and scientific methods of assessing the risks of biological control. Special attention should be paid to the specificity to the host. It is best to take into consideration the ratio of costs and benefits of biological control, which allows one to make a balanced decision on its implementation (Kenis et al., 2017).

Advantages and disadvantages of biological control Biological control has certain advantages in comparison with other strategies against pests, especially chemical methods. They include safety for the environment, absence of residues, affordability, ease of use, duration of impact, and absence of development of resistance in the pests. The disadvantages of biological control include: – Low tempi of control. There is often a delay between the increase in the population of pests and that of the agent of biocontrol. If a population of pests is already at the level of economic damage or higher, the only alternatives are pesticides. – Biological agents cannot eliminate the host completely, because this would cause their extinction. However, achieving complete elimination is possible through integration of biological control with other strategies against pests. – There is a possibility that a biological agent may try to feed on beneficial insects and plants. This problem is minimized through ac- curate selection of biological control agent. – There are concerns about release of foreign agents of control for they can bring new parasites which would affect the local community. 14 0 Chapter 3. Biological control of beetle pests of stored grain and field crops

– Methods of transport, maintenance and usage of a biological agent can be relatively difficult and in some cases expensive. – Biological control can be expansive compared chemical methods, which is related to the methods of study prior to the introduction of the strategy. – Badly prepared biological control can lead to rapid changes in the natural biodiversity (Kwenti, 2017).

General methods of using biological control Importation. The most broadly distributed classic method of biological control. It is used for the species of pests exploring a new environment, immigrants or introduced species, which have no natural enemies in the area. The method includes determination of the pest’s origin and determination of its natural enemies, which are potential agents of biocontrol. They are imported to a new place in the pest’s range with the aim of strengthening the species in new conditions and regulating the number of the targeted pest. Higher efficiency against the pests requires a strong capacity of the control agent for colonization. Maximum control is achieved if the agent is temporarily stable and can maintain its numbers even during brief absence of the target pest species (Mahr & Ridgway, 1993). Augmentation. Mass cultivation and release of natural enemies of the pest. Practically, there are several types of augmentation. Inductive release is one-time release of a large number of natural enemies in areas of different sizes. At the same time, the natural enemy does not have enough time to consolidate and does not reproduce. The method is used for one-time decrease in the numbers of the pest. Inoculating release is multi-time release of low numbers of the natural enemy over a certain period of time for its consolidation and distribution in the area of release. This method allows biological control of the pest’s population to be conducted over a long period of time. Maintenance. Using methods orientated towards protection, strengthening and support of existing natural populations of agents of biological control. Such natural enemies are already adapted to V. O. Martynov, V. V. Brygadyrenko 141 the environment and target pest, and their maintenance can be simple and economical. Such practice can include diversification of habitat for providing additional shelter or food to the natural enemy, using pesticides which are selective for target pests and cause minimum effect on natural enemies, provision of synthetic food additives, avoiding traditional agricultural methods which damage or eliminate natural enemies.

Agents of biological control

Predators Biological control of insects involves predators, parasites or pathogens such as viruses, bacteria and fungi. Predators are free living organisms, vertebrates or invertebrates which use pests as food and can consume them in large numbers. The efficiency of the control of the pest population by predators can vary depending on conditions in particular habitats. For example, in open areas, predation is up to 8 times higher than in the territory of an agrarian complex. In the areas with high grass stand, the level of predation is lower than in places with low grass. Spiders and Acari. Spiders hunt a variety of insects. After creating favourable living conditions for spiders, for example by mulching soil, one can increase their population by 60% (Riechert & Bishop, 1990; Jackson & Pollard, 1996). Some species of Acari, for example from Phytoseiidae family, are predators of nematodes and are able to consume eggs of Ascarididae from the soil (Lysek, 1963; Riechert & Bishop, 1990; Jackson & Pol- lard, 1996). Several species of Acari, such as Macrocheles muscaedomesticae (Scopoli, 1772), actively consume eggs and larvae of different flies which develop in the faeces of cattle (Safaa, 2014). Natural enemies of insect pests include the following Acari: Py- emotes tritici (LaGrèze-Fossat & Montagné, 1851), Blattisocius tarsalis (Berlese, 1918), Acarophenax mahundai (Steinkraus & Cross) and A. lacunatus (Cross & Krantz) (Bruce & LeCato, 1979; Haines, 14 2 Chapter 3. Biological control of beetle pests of stored grain and field crops

1981; Steinkraus & Cross, 1993; Faroni et al., 2000, 2001). The most studied species is P. tritici, but using this is limited because of harm for humans. P. ventricosus (Newport, 1850) is another much studied species, but its commercial use is limited because it can also bite humans, as do other species of the Pyemotidae family (Moser, 1975). Acari of the Acarophenacidae family are parasites of eggs of different pest insects. A. lacunatus affects Cryptolestes ferrugineus (Ste- phens, 1830), Rhyzopertha dominica (Fabricius, 1792) and Tribolium castaneum (Herbst, 1797), but cannot parasitize Oryzaephilus surinamensis (Linnaeus, 1758). In the studies by Oliveira et al. (2003) Acari significantly decreased populations of R. dominica by 61% and populations of T. castaneum by 53%. A. lacunatus significantly reduced the rapid tempi of growth of C. ferrugineus, R. dominica and T. castaneum populations. The range of hosts of A. lacunatus is not as broad as that of other parasitic Acari, agents of biological control of insects. For example, P. tritici attacks Plodia interpunctella (Hübner, 1813), Cadra cautella (Walker, 1863), Lasioderma serricorne (Fabricius, 1792) and even O. surinamensis (Bruce & LeCato, 1979). In the absence of the main host – С. cautella – B. tarsalis can parasitize T. castaneum. The ability to successfully parasitize several species is a good feature for potential use of a species as an agent of biological control of pests of grain crops (Oliveira et al., 2003).

Insects Successfully used classic agents of biological control are wasps of the Scoliidae family, such as Colpa sexmaculata (Fabricius, 1781). They are large wasps, the larvae of which parasitize larvae of the Scarabaeidae and Curculionidae families, while the imagoes feed on nectar. Parasites of nymphs and imagoes of many species of locusts are flies of the Sarcophaga genus, though their range of host preferences of is much wider. For example, Sarcophaga fuscicau- da (Böttcher, 1912) attacks adults of Rhynchophorus ferrugineus (Olivier, 1790) (Murphy & Briscoe, 1999). V. O. Martynov, V. V. Brygadyrenko 14 3

Representatives of the Tachinidae family are highly specialized agents of biological control, the larvae of which parasitize different species of Lepidoptera and Coleoptera. An example is Paratheresia menezesi (Townsend), which according to research, parasitizes 50% of populations of R. palmarum (Moura et al., 1993). Some earwig species of the Forficulidae family are active predators of insects. According to the data of Abraham et al. (1973), a single Chelisoches morio (Fabricius, 1775) consumes 5.3–8.5 of eggs and 4.2–6.7 larvae of R. ferrugineus daily. Predators broadly used against insect pests include the ware- house pirate bug Xylocoris flavipes (Reuter, 1884) and several other heteropterans of the Lyctocorinae subfamily (Reichmuth, 2000). These heteropterans are promising agents of control of Coleoptera and Lepidoptera in warehouses. They have strong ability to increase their number, even in conditions of prey shortage. However, X. flavipes is ineffective against Curculionidae species which hide in grains (Arbogast, 1975) and Teretriosoma nigrescens (Lewis) clown beetle (Richter et al., 1997). Another good example is Laelius pedatus (Say, 1836), a natural enemy of the varied carpet beetle (Anthrenus verbasci (Linnaeus, 1767)) and several species of Trogoderma (Klein & Beckage, 1990; Al-Kir- shi, 1998). In Northern Europe, for control of the housefly (Musca domestica (Linnaeus, 1758)), the predatory fly Hydrotaea aenescens (Wiedemann, 1830) is used commercially (Pirali-Kheirabadi, 2012). Promising agents of biological control of tsetse fly Glossina sp. (Wiedemann, 1830) are the velvet ants Mutilla glossinae (Turner) (Oluwafemi, 2008). Around 27 species of ants from Aphaenogaster, Iridomyrmex, Monomorium, Pheidole and Solenopsis genera hunt Acari, Haema- tobia irritans (Linnaeus, 1758) horn flies and different agricultural pests. In Louisiana (USA), Solenopsis invicta (Buren, 1972) was successfully used against Ixodidae ticks. However, this programme was not implemented on a large scale due to the ants’ danger to humans (Jemal & Hugh-Jones, 1993). 14 4 Chapter 3. Biological control of beetle pests of stored grain and field crops

An important role in the struggle against flies is played by dung beetles of the family Scarabaeidae which breed in cow faeces. The beetles Onthophagus ganelle and Euniticellus intermedius brought from Africa to Australia are used for the control of the Austra- lian bush fly (Musca vetustissima (Walker, 1849)) and buffalo fly (Haematobia exigua (de Meijere, 1906)), due to the fact that they compete with the fly larvae for food. Also, their rapid burial of dung reduces the range of places for nesting of the flies (Bornemissza, 1970; Pirali-Kheirabadi, 2012). For the control of the Colorado potato beetle (Leptinotarsa decemlineata (Say, 1824)), the spotted lady beetle (Coleomegilla maculate (De Geer, 1775)) can be used, which feeds on its eggs and larvae. Larvae of Coccinellidae are active predators which eat Aphids, Acari, small caterpillars and other insects (Acorn, 2007; Sing & Richard, 2008). For reducing the populations of C. cautella in warehouses, Bracon hebetor (Say, 1836) and Venturia canescens (Gravenhorst, 1829) parasitoids are used. Similarly, for reducing the population of O. surinamensis, the bethylid parasitoid wasp Cephalonmia tarsalis (Ashmead) is used. Other cosmopolite parasitoids of the wheat weevil are Lariophagus differendus (Forst) and Theocolax elegans (Westwood, 1874). Species used against P. interunctella are Tricho- gramma deion (Riley), which parasitizes eggs, and a larvae parasitoid B. hebetor, which significantly reduce the population of adults (Grieshop et al., 2006). Schmale et al. (2001) assessed the potential of several species of parasitoids of the bean weevil (Acanthoscelides obtectus (Say, 1831)). The most promising parasitoid for the control of A. obtectus is Di- narmus basalis (Rond). which has a long duration of reproduction and high number of offspring. A. calandrae turned out to be inap- propriate to be used as a control agent against this host. Heterospilus prosopidis (Viereck, 1910) had a shorter period of laying eggs than D. basalis, which led to lower life expectancy of its progeny. Amphibolus venator (Klug) is a predator of a number of insect pests of grain stores. It hunts Trogoderma granarium (Everts, 1898), T. castaneum, Corcyra cephalonica (Stainton, 1866), Latheticus oryzae V. O. Martynov, V. V. Brygadyrenko 14 5

(Waterhouse, 1880) and Alphitobius diaperinus (Panzer, 1797) (Pingale, 1954; Haines, 1991). Pingale (1954) mentioned that A. venator is an effective agent in the control of C. cautella and A. diaperinus. Nishi et al. (2004) in their study report that A. venator hunts larvae, pupae and adults of T. confusum, preferring the first. Predation is more significantly expressed at 30 °C than at 25 °C, because the optimum temperature for breeding and development of A. venator is 30.0–32.5 °C. Tempera- tures below 25 °C are unsuitable for the development of nymphs (Nishi & Takahashi, 2002). Females have a tendency towards consuming more prey compared to males because of the requirements of producing eggs. Another predator, Peregrinator biannulipes (Montrouzier & Signoret, 1861) is well known as a natural enemy of insect pests of grain stores. It hunts fungus moths such as Ephestia kuehniella (Zeller, 1879), P. interunctella, C. cephalonica, Pyralis farinalis (Linnaeus, 1758) and beetles, among which are T. castaneum, T. confusum, Stegobium paniceum (Linnaeus, 1758) and L. serricorne (Tawfik et al., 1983; Awadallah & Afifi, 1990). One of the most efficient parasitoids is a Pteromalidae wasp T. elegans, which attacks primary insect pests of grains, such as Si- tophilus spp., R. dominica, S. paniceum, Callosobruchus maculatus (Fabricius, 1775) and the Angoumois grain moth (Sitotroga cere- alella (Olivier, 1789)) (Cowan at al., 1881; Flinn et al., 1996; Flinn, 1998; Flinn & Hagstrum, 2001). However, T. elegans does not parasitize secondary pests, including Tribolium spp. and C. ferrugineus, the immature stages of which develop outside the kernel of wheat. In the study by Flinn et al. (2006), processing with this parasitoid led to significant decrease in the number of Sitophilus zeamais (Motschulsky, 1855), T. castaneum and C. ferrugineus by 78%, 94% and 70% respectively compared to the control group. Effective control of Indian meal moth (P. interpunctella) and other pyraloid moths was achieved using B. hebetor (Press et al., 1982; Cline et al., 1984; Cline & Press, 1990). Wasps of the Trichogramma genus, which are parasites of eggs of many lepidopteran pests, were also assessed for the control of P. interpunctella (Brower, 1990; Schol- ler & Flinn, 2000; Grieshop, 2005). 14 6 Chapter 3. Biological control of beetle pests of stored grain and field crops

Menon et al. (2002) studied the impact of A. calandrae which parasitizes R. dominica in wheat at different temperature ranges and host densities. It was determined that the efficiency of the parasitoid increases as the temperature rises. The highest number of infested larvae was observed at 35 °C and equaled 15 individuals in 24 h. At 20 °C, this parameter decreased to two larvae. At 38 °C, the death rate of the parasites significantly increases. Adult females of A. calandrae can survive up to 75 days in the presence of hosts and presence of cellulose as an additional source of food (Chatterji, 1955). It was found that female parasitoids survive up to 40 days if they have constant provision of hosts. Also, the parasitoid is a polyphage and is able to survive on different hosts such Sitophilus oryzae (Linnaeus, 1763) and S. zeamais (Williams & Floyd, 1971; Press et al., 1984). Therefore, the ability of A. calandrae to find and parasitize R. dominica in a wide range of temperatures makes it a good candidate for biological control of pests of grain storage in the areas with fre- quent fluctuations of temperature.

Pathogens

The conception of using pathogens The conception of using pathogens as agents of biological control appeared in the XIX century during the study of diseases of silkworm Bombyx mori (Linnaeus, 1758) (Tinsley, 1979). Already in 1880, Metchnikoff experimented with Metarhizium anisopliae ((Metch- nikoff) Sorokin, 1883), using it against pests of beet. As the role of pathogens in regulation of the insects was understood, the research focused on such biological problems as release of pathogens, their taxonomy, peculiarities of their life cycle, and further on comparative pathogenicity of different organisms (Mulligan et al., 1978; Baugher & Yendol, 1981), mechanisms of their transfer between hosts (An- dreadis & Hall, 1979) and on the ability of pathogens to maintain in the environment (Thompson & Scott, 1979). Also, researchers mentioned problems of development of methods of production, V. O. Martynov, V. V. Brygadyrenko 147 maintenance and transport of pathogens (Shapiro et al., 1981), their efficiency in field conditions (Ives & Cunningham, 1980) and problems of safety for humans and animals connected with usage of the agents (Harrap, 1978). The main attributes of a pathogen used as an agent of biological control should be high pathogenicity, high efficiency of the transfer, ability to survive inside the host, ease of production and maintenance in the laboratory (Anderson, 1982).

Fungi Many insects and other species of arthropods are vulnerable to infestation with entomopathogenic fungi. Currently, 750 species of entomopathogenic fungi are distinguished, which belong mostly to the Ascomycota and Zygomycota types. A distinctive peculiarity of fungi, in contrast to other pathogens is entrance into the host’s organism through the cuticle, not food. On contact of an insect with spores in the soil, air and on different surfaces, in the conditions of sufficient moisture, the spores germinate, forming hyphae, through the cuticle, mainly on bends of limbs where it is thinnest (Fathy, 2012). The tempi of the host’s death depends on the type of fungus and number of spores and usually occurs after 4–10 days. Also, fungi can release toxins (mycotoxin). After the host dies, the fungus produces thousands of new spores which scatter and continue their life cycle. Fungi are one of the most broadly used agents of biological pest control. The commonest species of entomopathogenic fungi are cosmopolite hyphomycetes M. anisopliae and Beauveria bassiana ((Bals.-Criv.) Vuill., 1912). In favourable conditions, they often cause natural outbreaks, affecting a broad range of insects and arachnids. In research on the development of these fungi, significant attempts were made to use them as control agents in agriculture and forestry of climatically mild regions. Because of their long survival in the soil, these fungi provide long term control, eliminating larvae and pupae of insects. Also, they are effective against mosquitoes, infesting them on different stages of development and causing their death 14 8 Chapter 3. Biological control of beetle pests of stored grain and field crops after 3–14 days (Quesada-Moraga et al., 2006; Leger, 2008; Ebrahim, 2015). The number of conidia released per host depends on the species of fungus, and species and size of host. For example, B. bassiana produces 10–200 times more conidia on adult Curculio caryae (Horn, 1873) than on M. anisopliae (Gottwald & Tedders, 1982). The most important abiotic factors which affect entomopathogenic fungi are temperature, moisture and ultraviolet radiation (Meikle et al., 2003). Other factors studied include moisture content and pH value (Lingg & Don- aldson, 1981), and soil structure (Groden & Lockwood, 1991). On the basis of B. bassiana, many products of Mycotrol ESO&WPO (BioWorks Inc.), Naturalis L (Troy BioSciences, Inc.) and Biosect® (KZ Chemicals) are available. As agents of biocontrol, oomycetes of Lagenidium giganteum (Schenk, 1859) are also used, which affect larvae of some species of mosquitoes. The fungus allows efficient control of the number of the mosquito Culex pipiens (Linnaeus, 1758) and as a result the spread of such diseases as filariasis and Rift Valley fever (Sur, 2001; Fathy, 2012). Another promising group of fungi which can cause natural outbreaks in insect populations is Entomophthora. Several genotypes of Entomophthora muscae ((Cohn) Fresen., 1856) were found, each being highly specific to a particular host (Jensen, 2011). Meyling & Eilenberg (2007) studied pathogenic Entomoph- thorales such as Pandora neoaphidis ((Remaudiere & Hennebert) Humber, 1989) and Neozygites fresenii ((Nowak.) Remaud. & Keller, 1980) as agents of biological control of aphids in England and the USA (Shah & Pell, 2003; Ekesi et al., 2005; Steinkraus, 2006). For insect control, Hirsutella thompsonii (Fisher), Lecanicillium lecanii (Zare & Gams), Nomuraea rileyi (Farl.), Coelomomyces spp. and Leptolegnia spp. (Fathy, 2012) have also been used. Paecilomyces formosoroseus and N. rileyi fungi can cause significant decrease in egg laying and emergence of adult pests of the Bruchidius genus. For the control of P. interpunctella, four species of fungi are used: B. bassiana, L. lecanii, M. anisopliae and Paecilomy- V. O. Martynov, V. V. Brygadyrenko 14 9 ces farinosus (Būda & Pečiulytė, 2008). According to Cuthbertson et al. (2005), Lecanicillium muscarium (R. Zare & W. Gams) is the most efficient agent of biological control used against some pests in orangeries and against the silverleaf whitefly (Bemisia tabaci (Gen- nadius, 1889)) (Schreiter et al., 1994; Faria & Wraight, 2001). There are reports ofB. bassiana killing Blissus leucopterus (Say, 1832) and Chrysoperla carnea (Stephens, 1836) (Ramoska & Todd, 1985; Donegan & Lighthart, 1989). It was demonstrated that adding 50 mg of conidia/kg leads to 50% decrease in the offspring, and 150 mg/kg – to 98% decrease, which is a maximum effect (Throne & Lord, 2004). Abd El-Aziz (2011) studied the impact of Bacillus thuringiensis (Berliner, 1915) and B. bassiana and three botanical extracts on three storage insect pests: Р. interunctella, E. cautella and E. kuehniella. Botanical extracts tested in combination with B. thuringiensis caused significant strengthening of pathogens and increase in the death rate in almost all cases (Sabbour, 2003). Būda & Pečiulytė (2008) tested the impact of four species of fungi (B. bassiana, L. lecanii, M. anisopliae and P. farinosus) on adult Indian meal moths (P. interpunctella) and demonstrated that all isolates were pathogenic, despite different levels of effectiveness. Gindin et al. (2006) studied the sensitivity of the snout beetle R. ferrugineus to the entomopathogenic fungi, M. anisopliae and B. bassiana. It was determined that the strains of the former were more virulent than the latter, leading to death of 100% of larvae over 6–7 days. Total percentage of death of eggs and hatched larvae equaled 80–82% compared to 34% in the control. The preparations were tested in the forms of powder and suspension. Using dry composition, 100% death of the adults was achieved after 2–3 weeks, and using suspension – after 4–5 weeks. The impact of the preparations reduced the life expectancy, the period of egg laying and caused a three-times decrease in the rate of hatching. Adane et al. (1996) studied the action of several isolates of B. bassiana against adult S. zeamais. The virulence of the studied isolates varied, the death rate of S. zeamais equaled 37–100%. Also, the 150 Chapter 3. Biological control of beetle pests of stored grain and field crops mean lethal time for different isolates was determined, the minimum value of which equaled 3 days. Lord (2001) studied the impact of B. bassiana on the snout beetle O. surinamensis and its control agent C. tarsalis. Processing the larvae of the snout beetle with fungi led to progeny of the wasp dying on all infested hosts two days after laying eggs and egg laying finishing on the fourth day after the processing. Decrease in egg laying is related to death of the host. It was determined that despite sensitivity to the fungus, the frequency of C. tarsalis visiting the storage site did not change, even when conidia were kept at 500 mg/kg of wheat, this being determined visually. The wasps are unable to find infested hosts and avoid lethal concentration of conidia on the surface of grains. The impact of B. bassiana at concentration of 100 mg/kg of wheat on the female C. tarsalis led to death of 53% of them in 3 hours. Male C. tarsalis proved less sensitive to the fungus. Khashaveh et al. (2011) studied the pathogenicity of M. anisopliae for T. granarium. A series of 105 to 109 conidia/ml suspensions were used in the experiment. All isolates were pathogenic for the adults and larvae, the death rate increased following increase in conidia concentration and significantly varied depending on the isolate. The overall death rate after 10 days was 21% to 84%, though one of the isolates demonstrated 100% death rate at maximum concentration. The study demonstrated that M. anisopliae is an acceptable agent of biological control of T. granarium, though additional experiments are required for increasing its efficiency. Most studies devoted to the interaction of parasitoid insects and fungi included fungi which do not attack parasitoids (Powell et al., 1986; Brobyn et al., 1988). In such cases, the parasitoid-fungi interaction is competitive rather than pathogenic. An example is the refuse of Encarsia formosa (Gahan, 1924) from the larvae of greenhouse whitefly with hyphae of mycelium of Asherhersonia aleyrodis (Webber) in their hemolymph (Fransen & Van Lenteren, 1993). Willers et al. (1982) reported on antifungal anal discharges of larvae of a Lepidoptera endoparasite, Pimpla turionellae (Linnaeus, 1758), which inhibit growth of B. bassiana. V. O. Martynov, V. V. Brygadyrenko 151

Bacteria The most important entomopathogenic bacteria belong to the Bacillus genus. One of the most broadly used agents of biological insect control is B. thuringiensis. After the bacteria enter the host, they release enterotoxins, which leads to the death of insect. For the control, different subspecies of B. thuringiensis are used: B. t. var. israelensis infests larvae of mosquitoes, black flies and related Dip- tera, B. t. var. kurstaki and B. t. var. aizawai – larvae of lepidopterans, B. t. var. tenebrionis – imagoes and larvae of beetles, B. t. var. japonensis – beetles which live in the soil (Fathy, 2012). B. t. var. kurstaki is able to control Acari actively compared to other subspecies (Hassanain et al., 1997). Usually, B. thuringiensis does not have a negative effect on parasitoids. Accordingly, Venturia canescens (Gravenhorst, 1829) contributes to distribution of this agent among E. kuehniella. However, B. thuringiensis causes pathogenic effect on a carrier parasitoid of Bracon brevicornis (Wesmael) (Schöller, 1998). Such products based on B. thuringiensis as DiPel® 2X DF, VectoBac® WDG and HD703 are available. Over the recent years, in Ukraine, the following microbiological preparations were studied and brought into production: Fitosporin, Hetomik, Bitoksibatsylin, Lepidotsyd, Bakteredentsyn and others. An effective agent of control of larvae of Culex spp. and Anopheles spp. mosquitoes is B. sphaericus (Robert et al., 1997). Streptomyces avermitilis bacteria produce avermectin toxins which are highly efficient against insects and other invertebrates (Pirali-Kheirabadi, 2012). There are data on infestation of R. ferrugineus with Pseudomonas aeruginosa ((Schroeter) Migula, 1900), a pathogenic bacteria which causes death of the insect in 8 days (Banerjee & Dangar, 1995). Symondson & Liddell (1996) reports that Wolbachia bacteria can have a negative effect on agents of natural control. It was noted that small and thin cocoons occur on the parasitoid Microctonus ae- thiopoides (Loan, 1975) which parasitizes Hypera postica (Gyllen- hal, 1813) infested with the bacteria, which causes death of 90% of the pupae and prevents adults reproducing. Therefore, Wolbachia 152 Chapter 3. Biological control of beetle pests of stored grain and field crops change the physiological conditions of the snout beetle, making it unsuitable for the parasite and providing the snout beetle protection.

Viruses Several thousands of pathogenic viruses have been described, and the representatives of Entomopoxviridae, Reoviridae (Cypovi- ruses) and Baculoviridae have been successfully used as agents of biological control (Lacey & Kaya, 2007). The infection occurs after the virus enters into the organism with food, and the peculiarities of replication varies depending on family. The most commonly used group of viruses is Baculoviridae, due to their specificity and safety for vertebrates. The Baculoviridae family contains four genera: Alphabaculovirus (lepidopteran-specific nucleo poly- hedroviruses (NPVs)), Betabaculovirus (lepidopteran-specific GVs), Gammabaculovirus (hymenopteran-specific NPVs) and Deltabaculovi- rus (dipteran-specific NPVs) (Jehle et al., 2006; Szewczyk et al., 2011). Currently, 16 baculovirus-based biopesticides are available, mostly for the control of lepidopterans. For example, the baculovirus of the codling moth (Cydia pomonella Granulovirus) is effective against their caterpillars, Gemstar LC is a NPV of Heliothis/Helicov- erpa spp. such as corn earworm (H. zea (Boddie, 1850)) and cotton bollworm (H. armigera (Hübner, 1805)), Spod-X LC is a NPV of Spodoptera spp. such as small mottled willow moth (S. exigua (Hübner, 1808)), CYD-X and Virosoft CP4 – C. pomonella Granu- lovirus and CLVLC is a NPVof celery looper (Anagrapha falcipera (Kirby, 1837)) (Fathy, 2012). Gopinadhan et al. (1990) found a highly efficient cytoplasmic polyhedrovirus (CPV) which affects R. ferrugineus at all stages of development.

Protists Entomopathogenic protozoa are studied insufficiently. Such protozoa as Nosema, Theileria and Babesia are pathogenic for Acari and other anthropods. There are no examples of effective biological control using protists, though such protozoa as Onchocerca volvulus V. O. Martynov, V. V. Brygadyrenko 153

(Bickel, 1982) and Plasmodium spp. are able to indirectly control intermediary hosts. Theratromyxa weberi, predatory soil amoeba, swallowed by nematodes, eats the host in 24 hours as they travel through their body. Presumably, such amoebas have limited abili- ties for biocontrol, as they are significantly slower than nematodes. Other protists, such as Nosema locustae (Canning, 1953), are pathogenic for katydids and crickets, Nosema pyrausta – for the European corn borer (Ostrinia nubilalis (Hübner, 1796)), and Vairimorpha ne- catrix – for caterpillars of many lepidopterans (Olson et al., 2006). Assessment of Microsporidia as agents of biological control of Prostephanus truncatus (Horn, 1878) was conducted by Henning- Helbig (1994). Gregarines are an interesting object of studies. This group of parasitic protozoa is studied insufficiently and they presumably occur in practically all species of insects. There are no data on usage of gregarines as agents of biological control. However, it was determined that gregarines can significantly decrease the efficiency of biological control by parasitizing the agent. Accordingly, Yaman et al. (2012) found gregarines of the Mattesia (Naville, 1930) genus in Rhizophagus grandis (Gyllenhal, 1827), a predatory beetle of the Mo- notomidae family, an agent of the control of great spruce bark beetle (Dendroctonus micans (Kugelann, 1794)). This gregarine also parasitizes victims which can be a source of invasion, though it causes no notable effect on its vital activity. At the same time, it was rather pathogenic for Rh. grandis. Mattesia parasitize the fat body of the host, in which merogony and sporogony occurs, leading to lysis of the tissues. This contributes to increase in life expectancy of the host and prolongation of the breeding period, leading to decrease in its potential of this agent for biological control. Lord (2003, 2006) studied the pathogenicity of Mattesia oryzaephili neogregarines which infest some pests of grain stores, including O. surinamensis and C. ferrugineus, and also their parasitoids such as C. tarsalis and C. waterstoni Gahan. Among wasps, the pathogen is distributed only among females during feeding on the host. Life expectancy of an infested female of C. tarsalis after contact 154 Chapter 3. Biological control of beetle pests of stored grain and field crops with highly infested O. surinamensis is 20 days, which is two times lower compared to a healthy individual. Infested C. waterstoni females survive 36 days instead of 46. M. oryzaephili can be transmit- ted through corpses, faeces of wasps and egg laying.

Nematodes Entomopathogenic nematodes are another variant of biological control, which can be used in combination with parasitoids and other strategies of biological control. They have been successfully used for regulating a large number of pests, including ground pests and wood-destroying pests (Canhilal & Carner, 2006; Mwaitulo et al., 2011; Kega et al., 2013). For distribution of the nematodes, there are used pulverizing devices and solutions of the agents which contain adjuvants and humidifiers – surface-active substances, oils and gels, which significantly increase the efficiency of the pest control, because nematodes survive only few hours on leaves due to their quick drying out and sensitivity to ultraviolet radiation (Arthurs et al., 2016). This indicates that entomopathogenic nematodes can be efficient against a broader range of insect pests, when combined with chemical additives prolonging their survival above ground (Baur et al., 1997; Schroer et al., 2005; Ditoet al., 2016; Noosidum et al., 2016). Nematodes of the Steinernema and Heterorhabditis genera are effective agents of control of a broad range of insect pests, including drain flies, the German cockroach (Blattella germanica (Linnaeus, 1767)), the cat flea (Ctenocephalides felis (Bouché, 1835)), Pennisetia marginata (Harris, 1839), leaf miners, mole crickets, flea beetles, plume moths, dark-winged fungus gnats, pyralid moths, owlet moths and others (Smart, 1995; Lacey & Kaya, 2007). Nematodes enter into the host organism with food, through the cuticle and spiracles. They contain specific symbiotic bacteria which cause death of the host and contribute to its decomposition, providing food to nematodes. Young nematodes consume a mix- ture of bacteria and liquefied tissue of insects, develop and breed inside the host. When the resources of nutrients are exhausted, V. O. Martynov, V. V. Brygadyrenko 155 the new generation emerges outside in search of new hosts (Grewal et al., 2005). For Steinernema, the presence of Xenorhab- dus spp. bacteria is typical, for Heterorhabditis – Photorhabdus spp. (Fathy, 2012). Entomopathogenic nematodes usually cause a high level of mortality of experimental insects. According to Caroli et al. (1996), Steinernema sp. caused death of 100% of Galleria mellonella (Linnaeus, 1758), Spodoptera exigua (Hübner, 1808) and Agrotis ip- silon (Hufnagel, 1766). For Heterorhabditis sp., lower mortality has been observed. Less sensitive to nematodes were Ostrinia nubilalis (Hübner, 1796) and Tenebrio molitor (Linnaeus, 1758), mortality of which equaled 73–100% for O. nubilalis and 20–100% for T. molitor. Also, the tempi of nematodes’ entry into the organism of T. molitor significantly varied depending on the species. The highest value was observed for S. glaseri and S. feltiae, the lowest for Heterorhabditis sp. (Caroli et al., 1996). An efficient agent of control is S. carpocapsae, an entomopathogenic nematode which parasitizes and sterilizes Musca autumnalis (De Geer, 1776). Portman et al. (2016) studied the virulence of several species of entomopathogenic nematodes for Cephus cinctus (Norton, 1872), a wheat pest. It was determined that the most effective species is H. indica (Poinar et al., 1992), because the mortality of the pest equaled 100% after two days. High concentrations of S. feltiae (200 and 500 IJ/larva) also caused 100% mortality after three days. The least harmful species studied was S. kraussei; the maximum mortality it caused was 60%. The obtained results indicate that H. indica and S. feltiae can be efficient agents of biological control of C. cinctus. Entomopathogenic nematodes are in widespread commercial use against insects. Currently, S. carpocapsae, S. riobraus, S. feltiae, S. glaseri, H. bacteriophora, H. megidis and H. marelatus are available, and also some nematodic products, Spear® and Saf T-Shield® (Kaya & Koppenhöfer, 1996). S. carpocapsae and S. glaseri are effective against mosquitoes, fleas and Acari, and also nematodes of Teladorsagia spp. and Trichostrongylus spp. (Henderson et al., 1995; Zhioua et al., 1995; Samish et al., 1996; Kocan et al., 1998). 156 Chapter 3. Biological control of beetle pests of stored grain and field crops

Nematodes of the Aphelenchida order have a broad range of relations with many species of Coleoptera (Hunt, 1993). They can range from simple commensalism to strictly parasitic relations. A well known example is the relationship between Bursaphelenchus cocophilus ((Cobb) Goodey) and Rhynchophorus palmarum (Linnaeus, 1758) in Central and South America. Rhynchophorus sp. were also observed to have two species of nematodes of Praecocilenchus. P. rhaphidiophorus (Poinar) parasitizes R. bilineatus (Montrouzier, 1857), and P. ferruginophorus (Rao & Reddy) parasitizes R. ferrugineus. Nematodes of both species occur in the trachea, intestines and the fat body of infested larvae, and also the womb and haemocoel of infested adults. The distribution of the nematodes occurs through oviposition or the intestines. It was determined that nematodes reduce the sizes of ovaries and egg production, and decrease life expectancy of snout beetles (Murphy & Briscoe, 1999). Ramos-Rodriguez et al. (2007) studied the efficiency of S. riobrave, an entomopathogenic nematode, against the storage pests T. castaneum and P. interunctella. Usage of the pathogen reduced the survivability of T. castaneum at all stages of its development from 78% to 27%. The temperature (25 and 30 °C) and relative moisture (40–100%) caused no significant effect on the efficiency ofS. riobrave in that experiment. During the usage of the agent in field conditions onT. castaneum and P. interunctella, the total survivability of the first equalled 42% and the total survivability of the second was 27% compared to the control. The larval stages of both species of insects were most sensitive to S. riobrave, the death rate equaling 99% and 80% for P. interunctella and T. castaneum respectively. S. riobrave is a promising agent of biological control for insect pests of grain warehouses. As biocontrol agents, entomopathogenic nematodes are safe, cause no harm to humans, vertebrates and plants.

Biological control of the main pests of wheat

Curculionidae The studies of biological control of S. oryzae were conducted by Riudavets & Lucas (2000). The natural enemies of these pests are V. O. Martynov, V. V. Brygadyrenko 157 parasitic wasps of Anisopteromalus calandrae (Howard, 1881) and Lariophagus distinguendus (Förster, 1841), which infest the snout beetle. Their potential against S. zeamais and S. oryzae was demonstrated in a number of studies (Arbogast & Mullen, 1990; Wen et al., 1994; Ryoo et al., 1996). Biological control with usage of these parasites leads to decrease in the number of progeny of S. oryzae by 57%. The efficiency of the control can be increased up to 90% by grain polishing. The development period of L. differentendus on S. oryzae is 20 days at 25 °C (Ryo et al., 1991), and of A. calandrae on S. zeamais – 17 days (Smith, 1992). The success of the control depends on several factors: grain polishing, which contributes to reducing the density of the pests; grain type which can inhibit the development of the stout beetle; integrity of laid eggs, which helps the parasite to identify its host. Also, the parasites exhibit great preference for larval stages of the fourth age, which indirectly affects the identification of the host as well (Smith, 1993; Ryoo et al., 1996). Smith (1994) studied and modeled the biological control of S. zeamais using A. calandrae, a parasitoid. It was determined that the long period of life and long period of laying eggs of S. zeamais hinders the control of its population by the parasitoid, though it is able to reduce the tempi of the snout beetle’s population growth. The imitation of the effect of releasing different numbers of A. calandrae demonstrated that there is no need to release over 10 times more parasitoids than the target species, and that the proportion 1: 1 is really efficient. The period between the first and the second generations of parasitoids allows the populations of snout beetle larvae to avoid parasitism, which causes the subsequent surge in the number of the pest after two weeks. Therefore, effective control requires minimum breaks in the continuity of parasitism. For the high efficiency, the optimum temporal interval between the successive releases of the parasitoid should not exceed 9 days at 25 °C. Throne & Lord (2004) provide data on the biological control of sawtoothed grain beetle (O. surinamensis) using an entomopathogenic fungus. The usage of the fungus at 300 mg conidia/kg of wheat 158 Chapter 3. Biological control of beetle pests of stored grain and field crops

(ppm) caused death to 72% of the adult individuals (Lord, 2001). The processing in 317 ppm led to death of 91% of larvae and pupae (Searle & Doberski, 1984). It was demonstrated that B. bassiana effi- ciently controls other pests, including S. zeamais (Adane et al., 1996, Hidalgo et al., 1998), S. oryzae (DalBello et al., 2001, Padin et al., 2002), P. truncates (Bourassa et al., 2001; Meikle et al., 2001), A. obtectus (Ferron, 1977), R. dominica, C. ferrugineus and T. castaneum (Rice & Cogburn, 1999; Lord, 2001). The experiments were conducted in the conditions favourable for O. surinamensis at 30 ± 1 °C and 76% relative moisture. Searle and Doberski obtained 91% decrease in larvae and pupae in the conditions of the culture at 20 °C and 100 relative moisture. Optimum temperature range for the growth of the most isolates of B. bassiana is 25–28 °C (Fargues et al., 1997). Sheeba et al. (2001) tested B. bassiana against the rice weevil (S. oryzae). It was determined that the strain used in study in the dosage of 7,6 log conidia/ml exhibits high efficiency at 28 ± 2 °C and relative moisture 70 ± 5%, leading to 76% mortality after 25 days, and reduction in the production of offspring by 86%. These results support the results obtained by Searle & Doberski (1984), where 100% mortality was recorded 20 days after the inoculation at 100% relative moisture and 25 °C. Trdan et al. (2005) studied the efficiency of four species of entomopathogenic nematodes (S. feltiae, S. carpocapsae, H. bacteriophora, H. megidis) for the control of S. granarius. A suspension of nematodes in 5,000, 10,000 and 20,000 IJs/ml concentrations was used at temperatures of 15, 20 and 25 °C over a week. It was determined that the least effective was H. megidis, whereas the remaining species had no significant differences in mortality rate they caused. Mortality of the beetles was statistically significantly higher at 20 and 25 °C. It has been proven that the impact of the suspension concentration was less important for the biological activity of the studied agents. Suspension in 20,000 IJs/ml proportion was most effective for the control of S. granarius (63%). No statistically significant differences between the other concentrations were found, though they V. O. Martynov, V. V. Brygadyrenko 159 were efficient in comparison with the control processing. Usage of the highest concentration (20.000 IJs/ml) of suspension was not jus- tified, because higher costs for purchasing biological agents do not provide significantly higher efficiency compared to the two other concentrations. S. feltiae, S. carpocapsae and H. bacteriophora were the most efficient against the pests, accordingly, all three species can be recommended as agents of biological control. Laznik et al. (2010) studied the effectiveness of several strains of S. feltiae against S. oryzae. The results demonstrated that the studied strains were most pathogenic at 25 °C and at highest concentration of nematode suspension (2,000 IJ/imagoes), mortality equalled 42– 72%. Minimum pathogenicity from 6% to 11% was observed at low concentrations of suspension and temperature of 30 °C. The lowest value of LC50 (1,165 IJs/imagoes) was obtained after 8 day exposure at 25 °C, whereas the highest (2,533 IJs/imagoes) – at 30 °C. Low efficiency of suspensions of all nematode strains at 30 °C, not ex- ceeding 20%, can be explained by the inappropriateness of the temperature intervals for most species of entomopathogenic nematodes, which equals 20–26 °C. S. oryzae reproduces more actively at high temperatures from 25 to 35 °C. According to the results obtained, the efficiency of S. feltiae depends on the temperature and level of concentration. There is a number of studies on the efficiency of S. feltiae against other storage pests, such as S. granarius (Trdan et al., 2006), T. confusum (Athanassiou et al., 2007), T. molitor, T. castaneum, T. variabile, S. oryzae (Ramos-Rodriguez et al., 2006) and O. surinamensis (Svendsen & Steenberg, 2000; Schöller et al., 2006; Tóth, 2006).

Tenebrionidae Rahman et al. (2009) studied the influence of X. flavipes on several species of pest insects. X. flavipes is a cosmopolite predator which affects such storage pests as T. castaneum, T. confusum, Cryp- tolestes pusillus (Schönherr, 1817), R. dominica and T. granarium (Ahmed, 1991). Arbogast (1976) reports that the victims include 13 species of insects from other orders. 160 Chapter 3. Biological control of beetle pests of stored grain and field crops

It was determined that the efficiency of X. flavipes depends on the sizes and density of the victims. In an empty medium, the predator killed a low number of large larvae of T. confusum and T. castaneum, compared to the other species of prey and living forms. This is related to the fact that the larvae can actively avoid the predator’s attacks, thus resisting their suppression and elimination. However, in wheat, this advantage disappeared, significantly increasing the effectiveness of the control. Also, it was found that X. flavipes prefers to prey on C. pusillus, compared to T. confusum and T. castaneum. Also, the efficiency of the control depends on the sex of the predator, for the females kill far more victims than the males. It should be mentioned that using X. flavipes in programmes of biological pest control in grain warehouses requires operational studies and surveys in field conditions. Wakefield (2006) studied the adhesion and germination of the conidia of B. bassiana on O. surinamensis and T. confusum. Con- centrations of 1х108 conidia/ml were used in the study. It was determined that the attachment of entomopathogenic fungi to the cuticle of insects is passive and non-specific (Boucias et al., 1988). Adhesion of the spores occurs through hydrophobic interactions (Jeffs et al., 1999). The highest mortality was recorded for O. surinamensis. For T. confusum, this indicator was significantly lower. Mortality of the control group equaled less than 5%. A high death rate was recorded for O. surinamensis, but for S. granarius, significantly lower death rates were observed, and even lower death rates for T. confusum. The death rate in the control equaled less than 5% for all three species. Few differences between the efficiency of two different isolates of B. bassiana were observed for each particular species. It was determined that the number of conidia and their germination on O. surinamensis are at significantly higher level than onT. confusum. This is related to the fact that O. surinamensis have much more setae and have a higher level of moisture on the surface of the cuticle, providing favourable conditions for germination of conidia. It was demonstrated that cuticular hydrocarbon can contribute to the germination of conidia or inhibit it. Also, Tribolium species produce protective quinones which, perhaps, can inhibit the germination. V. O. Martynov, V. V. Brygadyrenko 161

The potential of B. bassiana as an agent of biological control was studied by Khashaveh et al. (2011). They performed an experiment on adult individuals of T. castaneum, S. granarius and O. surinamensis, using 0, 250, 500, 750 and 1000 mg/kg concentrations over 5, 10 and 15 days at 24 ± 2 °C and 50 ± 5% moisture. In all experiments, the death rate increased as the dose and duration of exposure were increased. The highest values were observed after exposure to 1000 mg/kg concentration over 15 days and equalled 88%, 78% and 65% for S. granarius, O. surinamensis and T. castaneum respectively. Also, as the concentration increased, the production of offspring decreased. Michalaki et al. (2016) studied the effectiveness of M. anisopliae for biological control of T. confusum. Three dosages were tested – 8 · 106, 8 · 108 and 8 · 1010 conidia/kg of wheat or flour respectively. It was determined that at maximum concentration, the effectiveness of this form of control equaled 75%. Arthur (2000) studied the impact of diatomaceous earth on T. castaneum and T. confusum. The death rate of both species after the initial impact was minimum at 22 °C, but increased as the temperature and the interval of exposure increased, and decreased as the relative humidity increased. The highest mortality (100%) for both species was observed at 32 °C and 40% relative humidity. White & Loschiavo (1989) placed T. confusum in diatomaceous earth (0.72 mg/cm2) for 6 h, at 25 °C and 50% relative humidity, which led to absolute mortality after 3 days. McLaughllin (1994) placed S. granarius and S. oryzae, for 30 h at 25 °C and 56% RH in alumin- ium pots processed with diamaceous earth in the amount of 2 g/m2 and assessed the mortality during 6 days. In general, Tribolium spp. seem less sensitive to the inert dust and diatomaceous earth than other species of beetle pests of the stored grain (Korunic, 1998).

Competition in biological control Competition is an important element of biological control, which covers both target species of pests and the agents of control. Intraspecies and interspecies interactions include competition for 162 Chapter 3. Biological control of beetle pests of stored grain and field crops general resources such as food and space, and also predation and cannibalism (Park, 1962; Park et al., 1965, Stevens & Mertz, 1985). Introduction of agents of biological control significantly affects the competition of target species, causing changes in the structure of the community. A classic example of change in the outcome of competition between two species as a result of pathogenic invasion is the experimental study conducted by Park (1948) and Anderson et al. (1986). The study analyzed the competition between two species of flour beetle (T. confusum and T. castaneum) under the impact of Adelina tribolii (Bhatia, 1937), a species of Apicomplexa. In the absence of the parasite in a mixed population, the stable domination of T. confusum was observed, but after the pathogen was introduced, the domination disappeared. This is related to the fact that the parasite was pathogenic for the dominating competitor. Accordingly, the pathogen reduces the competitiveness of T. confusum, allowing a weak competitor to remain (Hudson, 1998). The impact of the rat tapeworm (Hymenolepis diminuta (Rudol- phi, 1819)) on the competition between two species of Tribolium was studied by Yan et al., (1998). Cestode parasitism causes a negative impact on the host`s reproductive success which affects the co- efficient of the host’s competition (Yan 1997). T. castaneum is usually a dominating competitor of T. confusum, for it is usually the most active among the species (Park, 1948, 1957; Leslie et al., 1968). Ac- cording to Craig (1986), while living together, adult species of T. castaneum mostly feed on eggs of T. confusum, because they are large. Prediction of the results of competition between the two species is complicated due to the impact of demographic stochastic processes (fluctuations in birth and death rates) and genetic stochastic processes (genetic drift with low number of founders) (Park, 1948; Park et al., 1964; Dawson & Lerner, 1966; Dawson, 1970; Mertz et al., 1976; Goodnight & Craig, 1996). T. castaneum is more vulnerable than T. confusum to infestation with H. diminuta (Yan & Norman, 1995). Therefore, the probability and extent of the invasion for T. castaneum is much higher. Parasitic invasion leads to decrease in the fertility of females, V. O. Martynov, V. V. Brygadyrenko 163 predominance of male and increase in cannibalism of eggs by the adults (Yan & Stevens, 1995). Accordingly, T. castaneum should be at a disadvantage in competition with T. confusum in the presence of the parasite. The results of the research demonstrate that the invasion of the cestode affected the competition between the two species of Tribo- lium, affecting the average time of death and population density. The presence of the parasite significantly increased the competitiveness of T. castaneum, especially in the case of high initial density of T. confusum. Mortality of T. confusum was increased by the parasite, therefore the species loses the competition with higher probability. The parasitism significantly reduced the average size of the population of T. confusum in the culture of the two species. This indicates that the invasion of the tapeworm gave T. castaneum an advantage in the competition with T. confusum. In this parasite-host-competitor system, T. castaneum is more sensitive to cestodes and has more clearly manifested pathogenesis than T. confusum, therefore having an advantage in competition. However, changes in the behavior, which were induced by the parasite, can give T. castaneum an advantage. Alabi et al. (2008) studied cannibalism and predation among seven species of Tribolium. The most usual type of cannibalism is consumption of passive forms (eggs and pupae) by the active forms (imagoes and larvae). Cannibalism of eggs is the most probable, for the eggs are least mobile, do not have highly developed protection, and at the same time contain a full set of nutrients in a form conve- nient for metabolism. One of highly developed forms of cannibalism is the production of trophic eggs, the aim of which is to be food for progeny, which is one of the forms of parental care (Crespi, 1992; Perry & Roitberg, 2006). Though pupae are also relatively immobile, they are more developed, sclerotized and can protect themselves (Eisner & Eisner, 1992). The highest tempi of egg cannibalsim was demonstrated by the larvae – they consumed 10 times more eggs than the imagoes of T. confusum and T. destructor. Among the adults of T. confusum 164 Chapter 3. Biological control of beetle pests of stored grain and field crops and T. brevicornis, they in total consumed 40, while by contrast T. freemani, consumed only a single egg. Imagoes of T. brevicornis, T. confusum and T. destructor demonstrate a greater tendency towards cannibalism than their larvae stages, whereas one can say exact the opposite, regarding T. anaphe, T. freemani and T. madens. Pupa cannibalism among the larvae is relatively low, with a maximum of 9 for T. anaphe. Among the imagoes, T. castaneum, T. confusum and T. destructor consumed 20 or more pupae, which is 4–5 times more than their larval stages, which was not manifested in other species. Regarding predation on T. castaneum, T. anaphe and T. brevicornis consumed 20 or 30 eggs respectively, which is 2 and 3 times more than the larvae of other species. The most voracious among the species were T. confusum and T. destructor, which consumed 54 and 28 eggs respectively. Pupae of T. castaneum were attacked mostly by the imagoes, the biggest consumer of which was T. destructor, consuming 34 pupae on average, whereas other species consumed less than ten. Research on predation on T. confusum showed that larvae of T. anaphe consumed 47 eggs on average, which is over double the consumption by T. brevicornis and T. madens. Among the imagoes, T. castaneum was the greatest predator both on eggs and pupae of T. confusum. When pupae of T. brevicornis were offered as victims, they were not consumed by any species, and predation was not higher than 5% for the larvae and 10% for the imagoes. According to results of the research, three groups of cannibals and predator species were distinguished in relation to the extent of cannibalism and predation. The first group was represented by T. castaneum, T. confusum and T. destructor, the adults of which are more voracious in cannibalism and predation than the larvae. The second group, where the larvae tended to be more active than the imagoes, consisted of T. anaphe, T. freemani and T. madens. The last group included only T. brevicornis, which is explained by variability in voracity for different types of prey among the larvae and adults of this species. Interspecies competition between natural enemies of a particular host can be an important factor in using biological control, because V. O. Martynov, V. V. Brygadyrenko 165 it influences the size, structure and stability of the agents’ communities (Ehler, 1978; Morris et al., 1988; Hagvar, 1989; van Alebeek et al., 1993; Chow & Mackauer, 1986). Interspecies competition between parasitoids can reduce the level of total parasitism and regulate the populations of pests (Turnbull & Chant, 1961; Watt, 1965; Pschorn- Walcher, 1977; Ehler & Hall, 1982). However, a minimization of the competition between the parasitoids may occur as a result of differentiation of the niche and interspecies specificity to the host, which leads to a significant inhibiting effect (DeBach, 1966; Huffaker et al., 1976; Keller, 1984). Parasitoids of the Pteromalidae family, A. calandrae and T. elegans attack immature stages of several insect pests of storages. The potential for inhibiting the rice weevil (S. oryzae) in wheat by the wasps A. calandrae and T. elegans was demonstrated in a number of works (Press et al., 1984; Cline et al., 1985; Press, 1992). Competition between A. calandrae and T. elegans was studied by Wen & Brower (1995). On combined release of the parasitoids, the domination of A. calandrae was observed, which was found three times more often than T. elegans, which coincides with the reports of Wen et al. (1994). However, the frequency of occurrence of both T. elegans and A. calandrae was significantly reduced when both were present compared to when they were released separately. The inhibition of the populations by competition can be a result of negative interactions between immature stages or caused by some activities of adult groups which affect the survivability of the larvae (Chow & Mackauer, 1986). The results of the experiment demonstrate the presence of a significant direct competition between the two parasitoids, which leads to significant decrease of both species. It should be mentioned that suppression of the populations of the rice weevil using a combination of the two species has the same effect as using only A. calandrae.

Conclusion Currently, biological control is one of the most promising and safe methods against pests. A large number of effective control agents 166 Chapter 3. Biological control of beetle pests of stored grain and field crops has been found, including predators, parasites and pathogens. Bio- logical control is one of the few methods against pests which has a stable pattern and which requires no activities after the release and distribution of the agents. In long term perspective, biological control has many advantages, such as safety for the environment, economic feasibility, ease in use, duration of the effect, absence of development of resistance in the pest. The main problem of biocontrol is the impact of agents on non- target species. On introduction of the control agent, the main task of the control is determination of the specificity of the agent to the host, because the main problem is the impact of agents on non-target species, which leads to changes in the communities. Before released into the environment, a potential agent of biological control should undergo a broad range of tests and quarantine. In selection of agents of biological control of a certain pest, one should take into account a number of factors such as population index, threshold effect, pest’s localization, species composition and stages of development (Burkholder & Faustini, 1991). However, there is a number of problems in the implementation of biological control including their potential effect on the natural diversity and unwillingness of farmers to give up using chemical methods (Heong & Escalada, 1998; Sun & Peng, 2007). Nevertheless, the methods of biological control continue to develop, adopting new technolo- gies, enlarging the usage of natural enemies, finding new agents of control and modifying old agents, for more efficient control of pests (Goswami et al., 2010; Ma et al., 2015). V. O. Martynov, V. V. Brygadyrenko 167

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Chapter 4. Fodder value of Poaceae family species in the steppe zone of Ukraine

B. O. Baranovsky, L. O. Karmyzova, I. А. Ivanko Oles Honchar Dnipro National University Introduction 1 Poaceae is one of the largest families of vascular plants. It has about 10 thousand species and 700 genera. Members of the family are spread worldwide. They often participate as dominants and edificators in composition of vegetation cover in grassy types of the Earth vegetation. Members of Poaceae family hold an important position within other plants (food, feeding, medicinal, industrial), useful for mankind. Poaceae are among the ten most widely represented families in all areas of the world. Participation of Poaceae, as well as other monocotyledonous plants decreases with the distance from the most East to moderate, and to equatorial latitudes (Tolmachev, 1974). Within the territory of Ukraine Poaceae family includes 71 genera (of which only 4 genera in the cultural state) and 208 species; of which only 15 genera are in the cultural state (Determinant of Higher Plants of Ukraine, 1987). Long-term anthropogenic influence on the territory of the steppe of Ukraine has led to a significant transformation of native vegetation. Nowadays, there is a significant reduction in species and ceno- tic diversity of ecosystems, in most of which the members of Poaceae family (grasses) dominate.

1 Baranovsky B. O., Karmyzova L. O., & Ivanko I. А. (2019). Fodder value of Poaceae family species in the steppe zone of Ukraine. In: Current problems of agrarian industry in Ukraine. Accent Graphics Communications & Publishing, Vancouver, Canada. – P. 191–227. Doi:10.15421/511905 192 Chapter 4. Fodder value of Poaceae family species in the steppe zone of Ukraine

According to the Decree of the Cabinet of Ministers of Ukraine on conducting the state accounting and Cadastre of flora in accordance with Article 38 of the Law of Ukraine “On fauna” and to the rati- fication by Ukraine of the International Convention on Biological Diversity at the initiative of the Ministry of environment in 2000 initiated the establishment of the State Plants Cadastre of Ukraine. It should serve as a scientifically substantiated basis for effective preservation and reproduction of phyto-diversity, rational use of resources of various plant groups. Grasses usually play the leading participation in herbaceous vegetation types. Many of them are edificators of steppe, meadow, mountain-meadow, mountain-steppe and other types of herbaceous phytocenoses. Most grasses are characterized by a wide ecological amplitude and occurs in several, or in many plant formations (Prokudin, Vovk, Petrova, 1977, Koreliakova, 1977). Grasses can also be used in phytoindication (Didukh, 1994; Dubyna, 1993). The main food plants of mankind belong to grasses are soft wheat (Triticum aestivum), rice (Oryza sativa) and corn (Zea mays), as well as many other crops (Zhukovsky, 1964). Usage of grasses as a forage plant for domestic animals is no less important. Economic value of grasses is also various in many other respects. Many grasses are also used in ornamental horticulture as lawn plants (Golovach, 1963; Abramashvili, 1970, Mytsyk, 2005, Lisovets, 2000). Grasses are also used for fixing mobile sands, various kinds of embankments, mine dumps (Shine,1956). Some grass species containing aromatic substances used in per- fumery, food industry and medicine. In our country, the most fa- mous species Seneca grass (Hierochloe) and sweet vernal grass (An- thoxanthum) contain coumarin, which is used to flavor for different beverages. Industrial application of grasses are also very various. Grasses have some negative effect, but it is completely incompa- rable with their benefits. Among the grasses, there are many weeds of different crops which cause significant damage. Especially it is B. O. Baranovsky, L. O. Karmyzova, I. А. Ivanko 193 applicable to wheatgrass, bromegrass, loose silky bent, bristle grass, barnyard grass, annual hairgrass. Grasses of the ruderal complex are competitors of forest stands at the creation of forest plantations in steppe (Belgard, 1950). Grasses are specially important as fodder plants for domestic animals (Shennikov,1941, Larin, 1956). Members of Poaceae family are the main components of natural hayfields and pastures on the territory of the steppe zone of Ukraine, especially of different types meadows and steppes (Prokudin, Vovk., Petrova, 1977).

Materials and methods of the survey The objects of the survey were representatives of Poaceae family in flora of the Steppe zone of Ukraine. Floristic surveys were carried out using common botanical methods, herbarization and identification of plant species (Manual of Plants of Ukraine, 1965, Skvortsov, 1977), as well as special methods for aquatic flora study (Katanskaya, 1981). Species determination was carried out using “Manual of Plants of Ukraine” (1965), “Determinant of higher plants of Ukraine (1987), “Flora of the USSR” (1935–1965), “Flora of the European part of the USSR” (1974–1989), “Flora of Eastern Europe” (1996–2004) using microscopes MBS1 and Cytoval. The list of species was compiled by based on the own collections, study of the herbarium of the Department of geobotany, soil science and ecology of DSU, comprisal of flora of the Prisamarya 1986 and 1988; the list was presented in alphabetical order (in Lat- in alphabet). Ecological certification of species was based on the Ecomorphs classification of O. L. Belgard (1950), which is the first ecomorphic system of vascular plant (Baranovsky et al., 1978). Ecological certification of plants was compiled with the use of literature: “Manual of Plants of Ukraine” (1965), “Flora of the USSR” (1935–1965), “Flora of the European part of the USSR” (1974–1989), “Flora of Eastern Europe” (1996–2004), “Macrophytes – indicators of environmen- 194 Chapter 4. Fodder value of Poaceae family species in the steppe zone of Ukraine tal pollution,1993”, monographs Would. O. Baranovsky (2000), V. V. Tarasov (2005) and V. V. Kucherevsky (2004). To establishment of the species ranges the following literature was used: “Manual of Plants of Ukraine” (1965), “Flora of the USSR” (1935– 1965), “Flora of the European part of the USSR” (1974–1989), “Flora of Eastern Europe” (1996–2004), “Horology of the flora of Ukraine” (1986), works Y. D. Kleopov (1990) and A. І. Kuzmichev (1992). Latin species and genera names are given on the latest nomenclature floristic edition of S. Mosyakin and M. М. Fedoronchuk (Mo- syakin, Fedorochuk, 1999).

Results In the steppe zone (Northern subzone) Ukraine Poaceae family amounted 149 species (Table 4.1). Table 4.1. – Ecomorphic characteristics of members of Poaceae family Name of plant Tropho- Hygro- Helio- Ceno- No spesies morphs morphs morphs morphs 1 2 3 4 5 6 Aegilops cylindrica PsPt 1. OgMsTr MsX He Нost Ru Aeluropus littoralis 2. AlkTr Ms He Hal (Gouan) Parl. Agropyron 3. OgTr MsX He RuPs dasyanthum Ledeb. Аgropyron. lavrenkoa- 4. OgTr MsX He Ps num Prokud. Аgropyron pectinatum 5. MsTr X He St (M. Bieb.) P. Beauv. Аgropyron tanaiticum 6. MsTr X He Ps Nevski .

7. Agrostis canina L. OgTr Ms ScHe StSMnPs B. O. Baranovsky, L. O. Karmyzova, I. А. Ivanko 195

1 2 3 4 5 6 Аgrostis capillaris L. 8. OgTr Ms ScHe SilPr (A. tenuis Sibth.) 9. Аgrostis gigantea Roth MsTr Ms ScHe SilPr Agrostis maeotica Alk- 10. XMs He HalPs Klokov OgTr Аgrostis sabulicola 11. OgTr Ms He PrPs Klokov 12. Аgrostis stolonifera L. OgMsTr Hg ScHe PrPal Аgrostis vinealis 13. OgTr Ms ScHe StSMnPs Schreb. Аlopecurus aequalis 14. OgTr HgHel He PrPal Sobol. Аlopecurus arundina- 15. AlkMgTr HgMs He HalPalPr ceus Poir. Аlopecurus genicula- 16. OgTr HgMs He PalPr tus L. Alopecurus pratensis 17. MgTr HgMs He Pr L. Anisantha sterilis (L.) 18. MsTr MsX ScHe PrStRu Nevski Anisantha tectorum 19. OgMgTr MsX ScHe PsRu (L.) Nevski Anthoxantum odora- 20. OgTr Ms ScHe SilPr tum L. Apera spica-venti (L.) 21. OgTr XMs ScHe RuPs P. B e auv . Arrenatherum elatius 22. MsTr XMs ScHe SilPr (L.) J. et C. Presl. 23. Avena fatua L. MsTr MsX He Ru 24. Аvena persica Steud. MsTr MsX He Ru 25. Аvena sativa L. MsTr MsX He RuCul 196 Chapter 4. Fodder value of Poaceae family species in the steppe zone of Ukraine

1 2 3 4 5 6 Beckmania erucifor- 26. AlkMsTr HgMs ScHe PalPr mis (L.) Host Botryochloa ich- 27. OgTrСа MsX He PtrSt aemum (L.) Keng Brachipodium sylvati- 28. MgTr Ms Sc Sil cum (Huds.) P. Beauv. 29. Briza media L. MsTr HgMs ScHe SilPr Вromopsis erectus 30. MsTr MsX He StRu (Huds.) Fourr. Bromopsis inermis 31. OgMgTr XMs He RuPrSt (Leyss.) Holub Bromopsis riparia 32. OgTr MsX He PrSt (Rehmer) Holub. 33. Bromus arvensis L. MsTr XMs He Ru Bromus commutatus 34. MsTr XMs He Ru Schrad. Вromus japоnicus 35. MsTr MsX He StRu Thumb. Вromus hordeaceus L. 36. MsTr XMs ScHe Ru (B. mollis L.) 37. Вromus secalinus L. MsTr Ms ScHe Ru RuPs 38. Bromus squarrosus L. OgMgTr MsX ScHe St Calamagrostis canes- SilPr 39. MsTr MsHg ScHe cens (Weber) Roth Pal Calamagrostis epigei- 40. OgMsTr Ms ScHe PsSilPr os (L.) Roth Catabrosa aquatica 41. MsTr Hel He PrPal (L.) P. Beauv. Cenchrus longispinus 42. ОgTr MsX He PsRu (Hack) Fernala B. O. Baranovsky, L. O. Karmyzova, I. А. Ivanko 197

1 2 3 4 5 6 Cleistogenes bulgarica 43. MsTr X He StPt (Bornm.) Keng Сleistogenes squarrosa 44. OgTr MsX He RuPs (Trin.) Keng Crypsis aculeata (L.) 45. AlkMsTr HgMs He HalPr Aiton Сrypsis alopecuroides 46. (Piller. et Mitterp) AlkMsTr HgMs He HalPs Schrad. Сrypsis schoenoides HalPs 47. AlkOgTr Ms He (L.) Lam. Pr Cynodon dactylon (L.) 48. AlkMsTr XMs He HalPr Pers. 49. Dactylis glomerata L. OgMsTr Ms ScHe SilPr Deschampsia caespitosa 50. MsTr MsHg ScHe SilPr (L.) P. Beauv. Digitaria aegyptica 51. OgTr MsX ScHe Ru (Retz.) Willd. Digitaria ischaemum 52. OgMsTr MsX He Ru (Schreb.) Muehl. Digitaria sanguinalis 53. OgMsTr Ms He PsRu (L.) Scop. Echinochloa сrusgalli 54. OgMgTr MsHg He Ru (L.) P. Beauv. Elymus caninus (L.) L. 55. (Roegneria canina (L.) MgTr Ms HeSc Sil Nevski Еlytrigia elongata 56. AlkTr Ms He PrHal (Host) Nevski Elytrigia intermidia 57. OgMsTr MsX ScHe StPtPs (Host) Nevski 198 Chapter 4. Fodder value of Poaceae family species in the steppe zone of Ukraine

1 2 3 4 5 6 Elytrigia repens (L.) MsX– SilSt- 58. MsTr ScHe Nevski MsHg PrRu Elytrigia stipifolia 59. (Czern. ex Nevski) MsTr MsX He PtSt Nevski Elytrigia trichophora 60. MsTr XMs ScHe SilSt (Link) Nevski Eragrostis aegyptica 61. OgTr XMs He Ps (Willd.) Delile 62. Eragrostis minor Host OgMsTr MsX He PsRu Eragrostis pilosa (L.) 63. OgTr MsX He RuPtPs P. B e auv. Eragrostis suaveolens MsX- 64. OgTr He SilPrPs Becker ex Claus HgMs Eremopyrum triti- 65. cetum (P. Gaertn.) AlkMsTr X He HalRuSt Nevski Festuca arietina 66. OgTr MsX ScHe SilPs Klokov Festuca beckeri 67. OgTr X He StSilPs (Hack.) Trautv. Festuca cretaceae 68. OgTr XMs ScHe Cr T. Pop. et Proskor. Festuca gigantea (L.) 69. MgTr HgMs Sc Sil Vill. 70. Festuca ovina L. s. l. MsTr X ScHe SilPr Festuca pratensis 71. MsTr HgMs ScHe Pr Huds. Festuca 72. pseudodalmatica MgTr X He HalPtSt Krajina ex Domin B. O. Baranovsky, L. O. Karmyzova, I. А. Ivanko 199

1 2 3 4 5 6 Festuca pseudovina 73. AlkMsTr X He HalSt Hack ex Wiesb. Festuca regeliana Pavl. 74. (F. orientalis (Hack.) AlkTr MsHg He HalPr V. Krecz. et Borbas) 75. Festuca rubra L. s. l. MgTr MsX ScHe SilPr Festuca rupicola 76. MgTr XMs He PrSt Heuff. Festuca valesiaca 77. MgTr X He St Goud. s.l. Glyceria arundinacea 78. MsTr HgHel He PrPal Kunth Glyceria fluitans (L.) 79. MsTr Hel ScHe PalAq R. Borbas Glyceria maxima 80. MsTr HgHel He PalAq (C. Hartm.) Holmberg Glyceria notata Che- 81. MsTr Hg He PrPal vall. Helictotrichon pubes- 82. MsTr XMs He StPr cens (Huds.) Pilg. Helictotrichon schelli- 83. MsTr MsX ScHe SMnPtSt anum (Hack.) Kitag. Hіerochloë odorata 84. OgMsTr XMs ScHe SilStPr (L.) Hіerochloë repens 85. OgMsTr XMs ScHe PsStPr (Host) P. Beauv. 86. Hordeum jubatum L. MsTr MsX He RuCul Hordeum leporinum 87. MsTr MsX He Ru Link 88. Hordeum murinum L. MsTr MsX He RuPsSt 89. Koeleria brevis Steven CaOgTr MsX He StPt 200 Chapter 4. Fodder value of Poaceae family species in the steppe zone of Ukraine

1 2 3 4 5 6 Koeleria cristata (L.) 90. MgTr X He St Pers. Koeleria delavignei 91. AlkMsTr XMs He HalStPr Gern. ex Domin Koeleria glauca 92. OgTr XMs ScHe SilPs (Spreng.) DC. Koeleria lobata 93. (M. Bieb.) Roem. et CaOgTr X He StPt Schult. Koeleria sabuletorum 94. OgTr MsX He PsSt (Domin) Klokov Koeleria talievii Lav- 95. OgTr XMs ScHe Cr renko Leersia orizoides (L.) 96. OgMsTr HgMs ScHe PrPal Sw. Leymus ramosus 97. MsAlkTr MsX He RuStHal (Trin.) Tzvelev Leymus sabulosus 98. OgTr MsX He PsRu (M. Bieb.) Tzvelev 99. Lolium perenne L. MgTr XMs He RuPr 100. Lolium temulentum L. MsTr Ms He Ru 101. Melica altissima L. MsTr XMs ScHe SMn Melica chrysolepis 102. MsTr MsX He PsPt Klokov 103. Melica nutans L. MsTr Ms Sc Sil 104. Melica picta K. Koch CaMsTr XMs ScHe Sil Melica transsilvanica 105. CaMsTr MsX ScHe SMnSt Schur 106. Millium effusum L. MgTr Ms Sc Sil Millium vernale 107. OgMsTr MsX He PsPrStPt M. Bieb. B. O. Baranovsky, L. O. Karmyzova, I. А. Ivanko 201

1 2 3 4 5 6 Molinia caerulea (L.) 108. OgTr Hg ScHe SilPrPal Moench 109. Nardus stricta L. OgTr Hg He SilPalPr 110. Panicum capillare L. OgMsTr MsX He Ru 111. Phalaris canariensis L. MgTr Ms ScHe CulRu Phalaroides arundina- 112. MgTr MsHg ScHe PrPal cea (L.) Rausch. Pholiurus pannonicus StPr 113. AlkTr MsX He (Host) Trin. Hal Phleum phleoides (L.) 114. MsTr XMs He PrSt Karst. 115. Phleum pratense L. MgTr Ms He Pr Phragmites australis 116. MsTr Hel ScHe PalAq (Сav.) Trin. ex Steud. 117. Poa angustifolia L. MsMgTr MsX ScHe SilPrSt 118. Poa annua L. MsTr Ms HeSc RuSilPr 119. Poa bulbosa L. OgMsTr MsX He RuSilSt 120. Poa compressa L. OgMsTr MsX ScHe RuSt 121. Poa nemoralis L. MsTr XMs ScHe Sil 122. Poa palustris L. MsTr MsHg He PalPr 123. Poa pratensis L. MsTr Ms He Pr 124. Poa remota Forcelles MsTr HgMs ScHe Sil 125. Poa sterilis M. Bieb. OgMsTr MsX He SilPt Poa erythropoda 126. MsTr MsX He PtSt Klokov 127. Poa sylvicola Guss. MgTr HgMs HeSc Sil PalPr 128. Poa trivialis L. MsTr HgMs He Sil PalPr Puccinellia bilykiana 129. AlkTr Ms He HalPr Klokov Puccinellia distans Ru Hal- 130. AlkMsTr XMs He (Jacq.) Parl. Pr 202 Chapter 4. Fodder value of Poaceae family species in the steppe zone of Ukraine

1 2 3 4 5 6 Puccinellia gigantea 131. AlkTr Ms He PrHal (Grossh.) Grossh. Sсlerochloa dura (L.) 132. MsTr XMs He StRu P. B e auv . Scolochloa festucacea 133. MsTr Hg He Pal (Willd.) Link 134. Secale sylvestre Host OgTr MsX He StRuPs Setaria glauca (L.) 135. MsTr XMs He PsRu P. B e auv . Setaria verticillata 136. MgTr Ms ScHe Ru (L.) P. Beauv. Setaria viridis (L.) 137. OgMsTr XMs He PsRu P. B e auv . Sorgum halepense (L.) 138. MsTr MsX He Ru Pers. Stipa asperella Klokov 139. MsTr X He StPt et Ossycznjuk 140. Stipa capillata L. MsTr X He PtSt Stipa borysthenica 141. OgTr MsX ScHe StPs Klokov ex Prokudin Stipa dasyphylla 142. (Czern. et Lindem.) MsTr X He St Trautv. Stipa lessingiana Trin. 143. MsTr X He St et Rupr. 144. Stipa pennata L. OgMsTr X He St Stipa pulcherrima 145. MsTr MsX He PtrSt K. Koch 146. Stipa tirsa Steven MsMgTr X He St Stipa ucrainica 147. MsTr X He St P. Smirn. B. O. Baranovsky, L. O. Karmyzova, I. А. Ivanko 203

1 2 3 4 5 6 Tragus racemosus (L.) 148. OgTr X He PsRu All. 149. Zizania latifolia Stapf MsTr Hel He Aq Note: Trophomorphs: OgTr – oligotroph (plant of poor soils); MsTr – mesotoph (plant of soil having moderate fertility level); MgTr – megatroph (plant of soil having high fertility level); AlkTr – alcotroph (plant of saline soils). Heliomorphs: He (Helio- phyton) – heliophyte (sun-loving); Sc (Sciophyton) – sciophyte (shade-tolerant). Hygromorphs: He (Helophiton) – helophyte (air-water); Hg (Hygrophiton) – hygrophyte (moist habitats); Ms (Mesophiton) – mesophyte (mid-moisture habitats); X (Xerophiton) – xerophyte (dry habitats). Cenomorphs: Aq (Aqa- nt) – aquant (water); Pal (Paludosus) – paludant (marsh); Pr (Pratensis) – patant (meadow); Sil (Silvaticus) – silvant (forest); SMn (Margosilvaticus) – silvomargoant (marginal species); St (Stepposus) – stepant (steppe); Ps (Рsammophyton) – psammophyte (growing on sandy soils species); Pt (Petrophyton) – petrophyte (species growing on the rocky habitats); Ru (Ruderatus) – ruderant (weed); Hal (Halophyton) – halophyte (species tolerant to high salinity); Cu (Cultus) – culturant (cultural species). 204 Chapter 4. Fodder value of Poaceae family species in the steppe zone of Ukraine

Hygromorphs

40 number of species 30 %

0 Ms X Hg Hel

Figure 4.1. The hygromorphs of most species of the grassesof the steppe zone of Ukraine

Heliomorphs

100 90 80 70 60 50 number of species 40 % 30 20 10 0 He Sc He HeSc Sc He

Figure 4.2. The heliomorphes of most species of the grassesof the steppe zone of Ukraine B. O. Baranovsky, L. O. Karmyzova, I. А. Ivanko 205

Among the members of vascular plants, flora of Poaceae family, mesophytes and xerophytes prevail in equal amounts, respectively 44% (66 species) and 43% (64 species). Numbers of hygrophytes and halophytes were fewer (10% (15 species) and 3% (4 species), respectively (Fig. 4.1). Among heliomorphes, heliophytes were dominated – 95% (142 species), taking into account the double biomorph (Fig. 4.2). Among trophomorphs, mesotrophs – 57% (85 species), oligotrophs 23% (34 species), megatrophs 15% (23 species) and alkotrophs 5% (7 species) were dominated (Fig. 4.3). Among cenomorphs, members of Poaceae family prevailed: pratants – 23% (34 species), stepants – 19% (28 species) and ruder- ants – 17% (26 species). Fewer: halophytes – 11% (17 species), sylva- nts – 6% (9 species), psammophytes – 6% (9 species), paludants – 5% (7 species), petrants – 4% (6 species), sylvomargoants – 3% (4 species), aquants – 3% (4 species), culturants – 2% (3 species) (Fig. 4.4).

Trophomorphs

90 80 70 60 50 number of species 40 % 30

20 10

0 OgTr Ms Tr MgTr AlkTr

Figure 4.3. The trophomorphs of most species of the grasses of the steppe zone of Ukraine A number of species (some forest and steppe species) demonstrated strict adherence to certain plant formations or habitat types 206 Chapter 4. Fodder value of Poaceae family species in the steppe zone of Ukraine

(halophytes, psammophytes, petrophytes, paludants). (Prokudin, 1977, Kucherevsky, 2004). 113 species (76%) have fodder value among the Рoaceae family in the steppe zone (Table 4.1). Among poisonous plant species, only two members of the same genera – Аgropyron tanaiticum Nevski and Agropyron dasyanthum Ledeb. were found.

Aq SMn 3% Cenomorphs 3% Cu Cr 2% 1%

Pt Pr 4% 23% Pal 5%

Ps 6%

Sil 6%

St 19% Hal 11%

Ru 17% Figure 4.4. The cenomorphs of most species of the grasses of the steppe zone of Ukraine Table 4.2. – Range, adventive status and co-zoological characteristics of Poaceae family members Status Name of plant Status of No Range of ad- Value species rarity ventity 1 2 3 4 5 6 Aegilops cylindrica 1. EM Forage Нost B. O. Baranovsky, L. O. Karmyzova, I. А. Ivanko 207

1 2 3 4 5 6 Agropyron dasyan- 3. End Pois. thum Ledeb. Аgropyron. lavren- 4. E Forage koanum Prokud. Аgropyron pecti- 5. natum (M. Bieb.) EAs Forage P. B e auv. Аgropyron tanaiti- 6. End Pois. cum Nevski Forage, 7. Agrostis canina L. EAs Ornam. Аgrostis capillaris L. 8. EAs Forage (A. tenuis Sibth.) Аgrostis gigantea 9. COSM Forage Roth Agrostis maeotica 10. PONT Forage Klokov Аgrostis sabulicola 11. End Forage Klokov 12. Аgrostis stolonifera L. EAs Forage Аgrostis vinealis 13. E Forage Schreb. Аlopecurus aequalis 14. EAs Forage Sobol. Аlopecurus arundi- 15. EAs Forage naceus Poir. Аlopecurus genicula- 16. EAs Forage tus L. Alopecurus pratensis 17. EAs Forage L. Anisantha sterilis (L.) 18. MIT adv. Nevski 208 Chapter 4. Fodder value of Poaceae family species in the steppe zone of Ukraine

1 2 3 4 5 6 Anthoxantum odora- Med., 20. Hol. tum L. Forage Apera spica-venti (L.) 21. ESIB adv. Forage P. B e auv. Arrenatherum elatius 22. EM Forage (L.) J. et C. Presl. Forage, 23. Avena fatua L. IT adv. Ruder. Forage, 24. Аvena persica Steud. EAs adv. Ruder. 25. Аvena sativa L. EAs adv. Forage Beckmania erucifor- 26. EAs Forage mis (L.) Host Botryochloa ich- Forage, 27. EAsM aemum (L.) Keng Indust. Brachipodium EAs- 28. sylvaticum (Huds.) Forage NAfr P. B e auv. 29. Briza media L. EAs Forage Вromopsis erectus 30. EAs Forage (Huds.) Fourr. Bromopsis inermis Forage, 31. EAs (Leyss.) Holub Phytomel Bromopsis riparia 32. EAs Forage (Rehmer) Holub. Bromopsis taurica 33. End Sljussar. 34. Bromus arvensis L. EAs adv. Ruder. Bromus commutatus 35. EMIT adv. Ruder. Schrad. Вromus japоnicus Forage, 36. EAs Thumb. Ruder. B. O. Baranovsky, L. O. Karmyzova, I. А. Ivanko 209

1 2 3 4 5 6 Вromus hordeaceus 37. COSM L. (B. mollis L.) 38. Вromus secalinus L. Hol. adv. Forage, 39. Bromus squarrosus L. EAs adv. Ruder. Calamagrostis canes- 40. EAs Forage cens (Weber) Roth Calamagrostis epigei- 41. EAs Forage os (L.) Roth Catabrosa aquatica 42. EAs Forage (L.) P. Beauv. Cenchrus longispinus S-NA 43. adv. Ruder. (Hack) Fernala COSM Cleistogenes bulgari- 44. EAs Forage ca (Bornm.) Keng Сleistogenes squarro- 45. EAs Forage sa (Trin.) Keng Crypsis aculeata (L.) 46. EAs Forage Aiton Сrypsis alopecuroides 47. (Piller. et Mitterp) EAs Forage Schrad. Сrypsis schoenoides 48. EAs Forage (L.) Lam. Forage, Cynodon dactylon 49. TROP adv. Lawn., (L.) Pers. Ruder. 50. Dactylis glomerata L. EAsAf Forage Deschampsia caespi- 51. Hol. Forage tosa (L.) P. Beauv. Digitaria aegyptica As 52. adv. Ruder. (Retz.) Willd. EM 210 Chapter 4. Fodder value of Poaceae family species in the steppe zone of Ukraine

1 2 3 4 5 6 Digitaria ischaemum E Forage, 53. adv. (Schreb.) Muehl. COSM Ruder. Digitaria sanguinalis 54. As adv. Ruder. (L.) Scop. Echinochloa сrusgalli Forage, 55. Hol. adv. (L.) P. Beauv. Ruder. Elymus canius (L.) EAs- 56. L. (Roegneria canina Forage SIB (L.) Nevski Еlytrigia elongata 57. PONT Forage (Host) Nevski Elytrigia intermidia 58. EAs Forage (Host) Nevski Med., Elytrigia repens (L.) 59. COSM Forage, Nevski Ruder. Elytrigia stipifolia RBU (Not 60. (Czern. ex Nevski) E Forage evaluated) Nevski Elytrigia trichophora 61. EAs Forage (Link) Nevski Eragrostis aegyptica 62. EAsAfr Forage (Willd.) Delile E 63. Eragrostis minor Host adv. Forage COSM Eragrostis pilosa (L.) E 64. adv. Forage P. B e auv. COSM Eragrostis suaveolens PONT- 65. Forage Becker ex Claus IT Eremopyrum triti- 66. cetum (P. Gaertn.) COSM Forage Nevski B. O. Baranovsky, L. O. Karmyzova, I. А. Ivanko 211

1 2 3 4 5 6 Festuca arietina 67. End Forage Klokov Festuca beckeri 68. EAs Forage (Hack.) Trautv. Festuca cretaceae 69. End T. Pop. et Proskor. Festuca gigantea (L.) EAs- 70. Forage Vill. SIB 71. Festuca ovina L. s. l. E Forage Festuca pratensis EAs- 72. Forage Huds. SIB Festuca EAs- 73. pseudodalmatica Forage SIB-M Krajina ex Domin Festuca pseudovina EAs- 74. Forage Hack ex Wiesb. SIB-M Festuca regeliana Pavl. (F. orientalis 75. EAs Forage (Hack.) V. Krecz. et Borbas) 76. Festuca rubra L. s. l. Hol. Forage Festuca rupicola 77. EAs Forage Heuff. Festuca valesiaca 78. E Forage Goud. s. l. Glyceria arundinacea Edible., 79. E Kunth Forage Glyceri. fluitans (L.) 80. E Forage R. Borbas Glyceria maxima EAs- 81. (C. Hartm.) Holm- SIB berg 212 Chapter 4. Fodder value of Poaceae family species in the steppe zone of Ukraine

1 2 3 4 5 6 Glyceria notata Che- 82. EAs Forage vall. Helictotrichon pubes- 83. EAs Forage cens (Huds.) Pilg. Helictotrichon schelli- 84. EAsNA anum (Hack.) Kitag. Hіerochloë odorata Med., 85. EAs (L.) Ruder. Hіerochloë repens 86. End (Host) P. Beauv. NA Decorat., 87. Hordeum jubatum L. adv. Hol. Ruder. Hordeum leporinum M 88. adv. Ruder. Link EMIT Koeleria brevis Ste- 89. E Forage ven Koeleria cristata (L.) 90. Hol. Forage Pers. Koeleria delavignei 91. EAs Forage Gern. ex Domin Koeleria glauca 92. ESIB (Spreng.) DC. Koeleria lobata E 93. (M. Bieb.) Roem. et Forage End Schult. Koeleria sabuletorum 94. EAs Forage (Domin) Klokov 95. Koeleria talievii Lavr. End Leersia orizoides (L.) 96. Hol. Sw. Leymus ramosus Forage, 97. EAs (Trin.) Tzvel. Ruder B. O. Baranovsky, L. O. Karmyzova, I. А. Ivanko 213

1 2 3 4 5 6 Leymus sabulosus 98. End (M. Bieb.) Tzvelev Forage, 99. Lolium perenne L. Hol. Decorat. Lolium temulentum M 100. adv. Ruder. L. COSM 101. Melica altissima L. EAs Forage Melica chrysolepis 102. E Forage Klokov 103. Melica nutans L. EAs Forage 104. Melica picta K. Koch E Forage Melica transsilvanica Forage, 105. EAs Schur Decorat. 106. Millium effusum L. EAs Millium vernale 107. MEAs M. Bieb. Molinia caerulea (L.) 108. EAs Forage Moench EAs- 109. Nardus stricta L. Forage NAfr 110. Panicum capillare L. NA adv. Ruder. Phalaris canariensis M 111. adv. Forage L. COSM Forage, Phalaroides arundi- Decorat., 112. EAs nacea (L.) Rausch. Water- protect. Pholiurus pannonicus 113. E Forage (Host) Trin. Phleum phleoides (L.) 114. EAs Forage Karst. 214 Chapter 4. Fodder value of Poaceae family species in the steppe zone of Ukraine

1 2 3 4 5 6 115. Phleum pratense L. EAs Forage Edible., Forage, Phragmites australis Decorat., 116. Hol. (Сav.) Trin. ex Steud. Water- protect., Indust. 117. Poa angustifolia L. EAs Forage 118. Poa annua L. Hol. Forage 119. Poa bulbosa L. MAs Forage Forage, 120. Poa compressa L. COSM Ruder. 121. Poa nemoralis L. EAs Forage 122. Poa palustris L. Hol. Forage Forage, 123. Poa pratensis L. Hol. Decorat. 124. Poa remota Forcelles ESIB Forage 125. Poa sterilis M. Bieb. E Poa stepposa (Kryl.) 126. PONT Roshev. 127. Poa sylvicola Guss. M Forage 128. Poa trivialis L. EAs Forage Puccinellia bilykiana 129. E Forage Klokov Puccinellia distans 130. EAs Forage (Jacq.) Parl. Puccinellia gigantea 131. E Forage (Grossh.) Grossh. Sсlerochloa dura (L.) 132. MIT adv. P. B e auv. B. O. Baranovsky, L. O. Karmyzova, I. А. Ivanko 215

1 2 3 4 5 6 Scolochloa festucacea 133. Hol. Forage (Willd.) Link 134. Secale sylvestre Host EAsAfr Ruder. Setaria glauca (L.) Edible., 135. As adv. P. B e auv. Ruder. Setaria verticillata 136. As adv. Ruder. (L.) P. Beauv. Setaria viridis (L.) Edible., 137. MIT adv. P. B e auv. Ruder. Sorghum halepense Forage, 138. M adv. (L.) Pers. Ruder. Stipa asperella Klok- RBU (Not Forage, 139. End ov et Ossycznjuk evaluated) Decorat. Stipa borysthenica RBU Forage, 140. EAs Klokov ex Prokudin (Vulnerable) Decorat. RBU (Not 141. Stipa capillata L. EAs Forage evaluated Stipa dasyphylla RBU PONT- Forage, 142. (Czern. et Lindem.) (Vulnerable) SIB Decorat. Trautv. Stipa lessingiana RBU (Not PONT- Forage, 143. Trin. et Rupr. evaluated SIB Decorat. RBU Forage, 144. Stipa pennata L. EAs (Vulnerable) Decorat. Stipa pulcherrima RBU Forage, 145. EAsM K. Koch (Vulnerable) Decorat. RBU PONT- 146. Stipa tirsa Steven Forage (Vulnerable) SIB Stipa ucrainica RBU (Not Forage, 147. PONT P. Smirn. evaluated Decorat. Tragus racemosus 148. M adv. Ruder. (L.) All. 216 Chapter 4. Fodder value of Poaceae family species in the steppe zone of Ukraine

1 2 3 4 5 6 Med., As 149. Zizania latifolia Stapf adv. edible., EAs Forage Note: RBU – species included in the “Red Book of Ukraine” (with types of rarity). Area: As – Asian; E – European; M – Mediterranean; IT – Iranian-Turanian; End – Endemic; COSM – Cosmopolite; PONT – Pontic; Hol. – Holarctic; TROP – Tropical; SIB – Siberian; NA, North American; SA – South and Central American The range of most species of the grasses of the steppe zone of Ukraine is Eurasian – 46% (68 species); lower number of species are representatives of the European area – 12% (18 species), cosmopolites – 8% (12 species), holarctic species – 8% (12 species), endemic –7% (10 species), Mediterranean – 7% (11 species), Asian – 7%(10 species) and Pontic areas – 5% (8 species) (Fig. 4.5). Area

PONT As 5% 7% M 7%

End EA s 7% 46%

Hol 8%

COSM 8% E 12% Figure 4.5 The range of most species of the grasses of the steppe zone of Ukraine B. O. Baranovsky, L. O. Karmyzova, I. А. Ivanko 217

Flora of Poaceae family in the steppe zone of Ukraine includes 21% (31 species) of the adventive element (Protopopova, 1991; Ba- ranovski, 2016). Flora of Poaceae family in the steppe zone of Ukraine includes 126 species having forage value. Within the flora of the steppe zone of Ukraine, there are 10 species were listed in the Red book of Ukraine (with the relevant types of rarity): Еlytrigia elongata (Host) Nevski – invaluable (Photo 4.5), Stipa asperella Klokov et Ossycznjukn– invaluable, Stipa borysthenica Klokov ex Prokudin – vulnerable, Stipa capillata L. – invaluable (Photo 4.2), Stipa dasyphylla (Czern. et Lindem.) Trautv. – vulnerable, Stipa lessingiana Trin. et Rupr. – invaluable (Photo 4.1), Stipa pennata L. – vulnerable (Photo 4.3), Stipa pulcherrima K. Koch – vulnerable, Stipa tirsa Steven – vulnerable (Photo 4.4), Stipa ucrainica P. Smirn. – invaluable.

Photo 4.1. Stipa lessingiana Trin. et Rupr 218 Chapter 4. Fodder value of Poaceae family species in the steppe zone of Ukraine

Photo 4.2. Stipa capillata L.

Photo 4.3. Stipa pennata L. B. O. Baranovsky, L. O. Karmyzova, I. А. Ivanko 219

Photo 4.4. Stipa tirsa Steven

Photo 4.5. Еlytrigia elongata (Host) Nevski 220 Chapter 4. Fodder value of Poaceae family species in the steppe zone of Ukraine

Discussion Usage of grasses as fodder plants for domestic animals of natural hayfields and pastures, especially meadows and steppes of various types is of great importance for the steppe zone of Ukraine. For that purpose it was useful up to 90% species of this family. Systematic structure of grasses in the steppe zone of Ukraine is quite various. Ecological and phytocenotic composition of grasses in various plant formations of the steppe zone of Ukraine has its own specific features (Belgard, 1950, Prokudin, 1977). Grasses in floodplain and ravine forests growing on the territory of the steppe zone of Ukraine have heterogenic composition. Shade- tolerant grasses (scyo-mesophytes) were typical for deciduous forests. Of these, the most common were: Brachypodium sylvaticum, Festuca gigantea, Poa nemoralis, Milium effusum, etc. Phytocenotic role of grasses in forest formations usually is insignificant. In most cases, they act as components of grass cover and play a subordinate role in its composition. Only some grasses dominate in the grass cover, as Poa nemoralis in lighter and drier types of broad-leaved forests of the Steppe. The number of other grasses in shady broad-leaved forests grows in more illuminated positions as at the edges, forest table lands and forest glades, in sparse forest site. Grasses of pine forests on sandy terraces of the rivers do not all relate to specific forest types. Most of them have a wider ecological amplitude and is also occurred in other vegetation types, such as Agrostis vinealis, Deschampsia caespitosa, Festuca ovina, Molinia coerulea, etc. Here, in conditions of better, compared with deciduous forests, light conditions, the number of grasses is growing. Calamagrostis epigeios often dominated in forests on sandy terraces (pure pine forests, mixed pine forest, birch forests), especially on their southern slopes. A large number of grasses increases sharply in open areas of the forest such as table lands and edges, cuttings and secondary meadows. In these habitats, however, they are mainly represented not by forest species but by components of other vegetation types, as well B. O. Baranovsky, L. O. Karmyzova, I. А. Ivanko 221 as by ruderal species (Belgard, 1950). In the context of the development of Ecomorph concept of O. L. Belgard, for species occurred on such biotopes it was selected the new cenomorph sylvomargoant – SMn (Baranowski, 2017). Grasses of plain-zonal steppe formations exhibit significant species diversity, have a different composition and a different interrelation of ecological groups. Xeromesophytes, xerophytes and mesophytes were dominated by the number of species; mesoxerophytes also were found in large numbers. Phytocoenotic role of grasses increases in these conditions. Fescue and many species of feather grass (in a broad sense) are of the main edificators of steppe plant communities. Additionally, the same (edificatory) role in the meadow steppes was belong to bush- grass (Calamagrostis epigeios), in the meadow and forb-fescue-feather grass steppes – narrow-leaved bluegrass (Poa angustifolia) and awn- less bromegrass (Bromopsis inermis), in the forb-fescue-feather grass steppes – meadow brome (Bromopsis riparia), in the fescue-feather grass steppes – meadow brome, in the desert steppes – Wheatgrass comb (Agropyron pectinatum). The abovementioned and other grass species are part of many steppe associations and association groups as dominants, subdominants or as typical species. Composition and distribution pattern of these grasses also varied according to zonal, climatic and soil conditions. Grass species associated with hard rock exposures have a number of features inherent to steppe grasses. Within the steppe zone of Ukraine on hard rock exposures the same number of grass species as in the steppes was observed. Both have many common species, about 50%. There is a great similarity between them in the interrelation of ecological groups. On hard rock exposures, different species of comon steep grasses were widely represented: Festuca sulcata s. і., Koeleria cristata, Stipa caerulea, Agropyron pectinatum, as well as specific – Botryochloa ichaemum. They often dominated in the grass cover, especially in areas with of erosion process decaying. A large number of grass species was found in flora of sandy steppes on river terraces. 222 Chapter 4. Fodder value of Poaceae family species in the steppe zone of Ukraine

By the species composition, sand grasses are some similar to that of steppes (26% of the total species) and of hard rock exposures (24% of the total species), although they include a significant number of specific species – psammophytes, confined only to given edaphic variant. Ecologically, xeromesophytes are predominant group under such environmental conditions with a numerous species; also there is quite number of xerophytes occurred. The species composition of grasses and their participation in the composition of the vegetation cover of sandy steppes were largely depend on sand mobility and level of their overgrowth. On the sandy river terraces slightly overgrown by vegetation, xerophilic (Agropyron dasyanthum, A. tanaiticum) and mesophilic (Calamagrostis epigeios) long-root grasses were dominated. They act as dominants and co-dominants and form one – or low-species shrubs. Psammophyte grasses were typical for overgrown sands, among which the sod grass species were the most common: Agropyron lavrenkoanum, Festuca beckeri, Koeleria sabuletorum, Stipa borysthenica. They often participate as co-domi- nant in the vegetation cover. Meadow grasses were represented by a large number of species (Dmitrieva, 1982, Prokudin, 1977), among which mesophytes were predominated. Depending on the topography and level of soil moisture, interrelation of ecological groups of grasses was different. Grasses were mainly involved in the composition of meadow phytocenoses. Foxtail grass (Alopecurus pratensis), meadow fescue (Festuca pratensis), meadow grass (Poa pratensis) formations were common on both watershed and floodplain meadows. On the steppe meadows formation, Welsh fescue (Festuca valesiaca), brown bent (Аgrostis vinealis), narrow-leaved bluegrass (Poa angustifolia) the most commonly occurred. Wet meadows were formed with the formation of reed sweet grass (Glyceria maxima), floating grass (Glyceria fluitans), reedy canary grass (Phalaroides arundinacea), Beckman’s grass (Beckmania eruciformis), swamp meadow grass (Poa palustris) and other grass species. B. O. Baranovsky, L. O. Karmyzova, I. А. Ivanko 223

Floristically, the grasses of swamps and ponds were the most close to the meadow grasses, but they distinct from each other by the poorest species composition and predominance of hygrophytes (Afanasiev;1968). Grasses of swamps and ponds are Phragmites australis, Calamagrostis canescens, Glyceria maxima, Leersia oryzoides. Of all the grass species in the steppe zone of Ukraine, one of the most common in shallow water and flooded areas (Baranovsky, 2000) is a common reed grass (Phragmites australis) which was also used in very dry years as a forage crop. This is because its high vegetative mobility, generative ability, environmental plasticity, resistance to anthropogenic pollution. But in this regard, common reed grass has a negative economic value more than positive because (Macrophytes as indicators of changes of natural environment, 1993). Phytocenotic participation of common reed grass (Phragmites australis) was especially great in shallow areas of reservoirs and ponds (Baranovsky, 2000). It often forms single-species phytocenoses with large area and biomass (above 10 kg/m2) at depths from 0 to 2.0 m. Reed sweet grass (Glyceria maxima) and glyceria reed (Glyceria arundinacea) can form single-species formations. Other grass species are more often components of the vegetation cover in exces- sively moist habitats. In the vegetation composition of salt marshes, grasses were represented mainly by mesophytes. Among them there are many species also typical for meadow cenoses (up to 50%). Species of Puccinella genus (Chorology of flora, 1986, Belgard, 1950) were found among the dominants in the vegetation cover of saline pods, lowland wetlands and floodplain meadows, as well as on the meadows of the coastal strip. Flora of Poaceae family in the steppe zone of Ukraine includes 126 species that have forage value.

Conclusion Poaceae family in the flora of the steppe zone of Ukraine is represented by 149 species belonging to 56 genera. The largest number of species belongs to the genera: Poa (meadow grass), Festuca repre- 224 Chapter 4. Fodder value of Poaceae family species in the steppe zone of Ukraine sented by 12 species within the Steppe of Ukraine, and Stipa (stipa grass) – 7 species, but they are all rare. The result ecomorphic analysis of Poacea flora of Dnipropetro- vsk Oblast showed that hemicryptophytes were the most numerous (55.7%), and helophytes were less numerous (6.4%) among clima- morphes. Among hygromorphes, meso-xerophytes were most numerous (30%), and hygrophytes and meso-hygrophytes were lesser (5%). Heliophytes dominated among heliomorphes (64.3%), and scyophytes have the lowest proportion (1.4%). Majority of tropho- morphes was represented by mesotrophes (38.5%), and the smallest number have alco-megatrophes (0.7%). Among cenomorphes, the largest share in the Poaceae flora was present in stepants and pratants (17.7% and 25.5%, respectively). All this contribute to a wide approach to pasture use of steppe, meadow and marginal areas with the dominance of grasses in the steppe zone of Ukraine. Flora of Poaceae family in the steppe zone of Ukraine includes 126 species that have forage value. In the flora of grasses of steppe zone of Ukraine there are 3 species listed in the World Red List (Agropyron dasyanthum Ledeb. – northern wheat grass, Elytrigia stipifolia (Czern. ex Nevski) Nevski – Feathergrass-leaved wheatgrass, Stipa dasyphylla (Czern. et Lindem.) Trautv. – feather grass), 1 species included in the European Red List (Elytrigia stipifolia (Czern. ex Nevski) Nevski – Feathergrass-leaved wheatgrass) and 9 species are listed in the Red Book of Ukraine. All these species need to be protected, and the use of pastures with their participation should be limited. B. O. Baranovsky, L. O. Karmyzova, I. А. Ivanko 225

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Contents

N. M. Zazharska Chapter 1. Assessment of safety and quality of goat’s milk. . . . . 3

V. I. Rusynov, V. O. Martynov, T. M. Kolombar Chapter 2. Coleoptera pests of stored food supplies and field crops...... 34

V. O. Martynov, V. V. Brygadyrenko Chapter 3. Biological control of beetle pests of stored grain and field crops...... 134

B. O. Baranovsky, L. O. Karmyzova, I. А. Ivanko Chapter 4. Fodder value of Poaceae family species in the steppe zone of Ukraine ...... 191